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Volume 30 Number 2,

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1987

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ISSN 0029-8182

Oceanus

The International Magazine of Marine Science and Policy

Volume 30, Number 2, Summer 1987

Paul R. Ryan, Editor
James H. W. Main, Assistant Editor
Michelle K. Slowey, Editorial Assistant
T. M. Hawley, Spring Intern

Editorial Advisory Board

1930

Henry Charnock, Professor of Physical Oceanography, University of Southampton, England

Edward D. Goldberg, Professor of Chemistry, Scripps Institution of Oceanography

Gotthilf Hempel, Director of the Alfred Wegener Institute for Polar Research, West Germany

Charles D. Hollister, Dean of Graduate Studies, Woods Hole Oceanographic Institution

John Imbrie, Henry L. Doherty Professor of Oceanography, Brown University

John A. Knauss, Provost for Marine Affairs, University of Rhode Island

Arthur E. Maxwell, Director of the Institute for Geophysics, University of Texas

Timothy R. Parsons, Professor, Institute of Oceanography, University of British Columbia, Canada

Allan R. Robinson, Gordon McKay Professor of Geophysical Fluid Dynamics, Harvard University

David A. Ross, Chairman, Department of Geology and Geophysics, and Sea Grant Coordinator,

Woods Hole Oceanographic Institution

Published by Woods Hole Oceanographic Institution

Guy W. Nichols, Chairman, Board of Trustees
lames S. Coles, President of the Associates

John H. Steele, President of the Corporation
and Director of the Institution

The views expressed in Oceanus are those of the authors and do not
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COVER: A marine iguana foraging for algae on the bottom in Galapagos waters. Photo
by Edmund S. Hobson © National Geographic Society. BACK COVER: Marine iguana
colony onshore. Photo by Flip Schulke, Black Star © National Geographic Society.

Copyright © 1 987 by the Woods Hole Oceanographic Institution. Oceanus (ISSN 0029-
8182) is published in March, June, September, and December by the Woods Hole
Oceanographic Institution, 93 Water Street, Woods Hole, Massachusetts 02543. Sec-
ond-class postage paid at Falmouth, Massachusetts; Windsor, Ontario; and additional
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Oceanus

Woods Hole Oceanographic Institution
Woods Hole, Mass. 02543

Subscription correspondence, U.S. and Canada: All orders should be addressed to Oceanus

Subscriber Service Center, P.O. Box 6419, Syracuse, N.Y. 13217. Individual subscription rate: $22 a

year; Libraries and institutions, $50. Current copy price, $5.50—25% discount on current copy

orders for 5 or more; 40% discount to bookstores and newsstands. Please make checks payable to

Woods Hole Oceanographic Institution.

Subscribers outside the U.S. and Canada, please write: Oceanus, Cambridge University Press, the

Edinburgh Building, Shaftesbury Rd., Cambridge CB2 2RU, England. Individual subscription rate

£20 a year; Students, £17; Libraries and Institutions, £37. Single copy price, £9. Make checks

payable to Cambridge University Press.

When sending change of address, please include mailing label. Claims for missing numbers from

the U.S. and Canada will be honored within 3 months of publication; overseas, 5 months.

4
6

9
16

20
28
33
42
49
54
61
69

Foreword

by Leon Febres Cordero, President of Ecuador

President's Decree on Galapagos Marine Resources Reserve

A Promise to the Sea, and the Politics of the Decree

by Roque Sevilla

The Galapagos Marine Resources Reserve and Tourism Development

by lames M. Broadus

Two Legal Opinions on the Galapagos Marine Reserve
16 — Ecuadorian Law

by Efrain Perez Comacho
17 — International Issues

by Kilaparti Ramakrishna

Diving in the Galapagos

by Codfrey Merlen

The Fishes of the Galapagos Islands

by lohn E. McCosker

Marine Biological Research in the Galapagos: Past, Present, and Future

by Henk W. Kasteleijn

Negative Effects of the 1982-83 El Nino on Galapagos Marine Life

by Gary R. Robinson

Sperm Whale Behavior on the Galapagos Grounds

by Hal Whitehead

Marine Iguanas: Living on the Ocean Margin

by Andrew Laurie

The Urvina Bay Uplift: A Dry Trek Through a Galapagos Coral Reef

by Mitchell W. Colgan, and David L. Malmquist

A Search for Unique Drugs in the Galapagos Underwater Environment

by Shirley A. Pompon;, and Susan van Hoek

72 The Voyage of the Beagle, Chapter 17 (edited)

by Charles Darwin

79 Darwin in the Galapagos: Three Myths

by Frank /. Sulloway

86 Whalers, Whales, and Tortoises

by Bruce E . Epler

93 Galapagos Tales

by Paul R. Ryan

1 00 Book Reviews

Copyright © 1987 by the Woods Hole Oceanographic Institution. Oceanus (ISSN 0029-
8182) is published in March, June, September, and December by the Woods Hole
Oceanographic Institution, 93 Water Street, Woods Hole, Massachusetts 02543. Sec-
ond-class postage paid at Falmouth, Massachusetts; Windsor, Ontario; and additional
mailing points. POSTMASTER: Send address changes to Oceanus Subscriber Service
Center, P.O. Box 6419, Syracuse, N.Y. 13217.

92C

91°

REPUBLICA DEL ECUADOR
ARCHIPIELAGO DE COLON

\ (PROVINCIA INSULAR DE GALA'PAGOS)

DARWIN (CULPEPPER) IS

168 '

WOLF (WENIMAN) IS. »

Internal
Waters

50,000
(sq kilometers)

Redondo Rock

FERNANDINA
(NARBOROUGH)IS

1494

ISABELA
(ALBEMARLE)IS.

A Puerto Villamil/
\ \

Pmzon Beagle Is

(Duncan) Is

Guy Fawkes Is

Grossman Is
Tortuga
(Brattle) Is

Puerto Ayora

SANTIAGO

(JAMES)

906

PINTA (ABINGDON) IS.

777 \

(BINDLOE) IS.

343

GENOVESA

(TOWER) IS.

Seymour Is

Baltra Is A

SANTA JCRUZ

^(INDEFATIGABLE) IS. /

Plaza Is

Acode

Puerto
Velosco Ibarra

FLOREANA
(CHARLES)

640

Elevation in meters

• Airport

• Population Center

ESPANOLA
(HOOD) IS

\ 199

SAN CRISTOBAL
(CHATHAM) IS.

716

20 40

Kilometers

90C

IOC

89C

90° 2° 89° 1°

The Galapagos Marine Resources Reserve — established by Ecuadorian Presidential Decree on May 13, 1986 — defined as
including the water column, seabed, and marine subsoil of the interior of the Galapagos Archipelago — (he area within the
baselines drawn around the outer points of the islands — and within a band of 15 nautical miles surrounding these baselines. The
total of 19 islands and 42 islets are located in the Pacific Ocean, 600 miles west of Ecuador, directly on the equator. The land
area is 7,882 square kilometers with a coastline of 1,350 kilometers. The capital of the Galapagos is Puerto Baquerizo Moreno,
on San Cristobal Island. Puerto Ayora is the largest city, with an approximate population of 5,000 out of a total Galapagos
population of 8,000 to 10,000 (only 4 islands are inhabited). Elevations and depths are in meters. Boundary lines are an
approximate representation. The internal waters of the reserve amount to some 50,000 square kilometers, with an additional
20,000 square kilometers representing the approximate figure for the buffer zone — a total of about 70,000 square kilometers.

El Ecuador ha sido, es
y seri Pals Amaionico

PRESIDENCIA DE LA REPUBLICA

March 2, 1987.

To Oceanus magazine in celebration of its special issue on the new Galapagos Marine Reserve.

The territory of Ecuador is divided by the Equator and is like two outstretched hands greeting both
the Northern and Southern Hemispheres.

Ecuador is also divided vertically by the majestic Andes cordillera that towers more than
18,000 feet, with snow-crowned peaks only a few hundred kilometers away from the Pacific
Ocean and the Amazon Basin. These factors combine into privileged situations. Our country has all
the climates of the Earth without their extreme rigors. Its ecosystems are unique: Ecuadorian flora
and fauna are extraordinarily rich and diverse and can still be found in their almost unaltered
natural habitats.

Among all these wonders, is the Galapagos Islands, the "Enchanted Islands," cast upon the
ocean like a handful of pearls. It was on these islands that Charles Darwin conceived his famous
theory of the evolution of species by means of natural selection, perhaps because animal species,
long extinct in other parts of the world, still live among the Galapagos volcanic lava and white, soft
beaches as reminders of centuries past.

The Ecuadorian Government has taken appropriate measures to preserve, protect, and
conserve the resources of the Galapagos Islands, especially the water column, the seabed, and the
submarine subsoil of the archipelago's interior seas.

The Ecuadorian Government therefore sponsors and praises international efforts that
contribute to these objectives. One must not forget that even though Ecuador maintains
sovereignty over the Galapagos Islands and their surrounding waters, these islands also belong to
the cultural heritage of all humankind.

CORDERO R

PRESIDENTE CONSTITUC IONAL DE LA REPUBLICA
DEL ECUADOR.

The Galapagos

Marine Resources Reserve Decree

The Official Register of Ecuador— Number 434— May 13, 1986,
pp. 28-29

No. 1810-A

Leon Febres-Cordero Ribadeneyra
Constitutional President of the Republic

Whereas

It is the duty of the National Government to protect those natural areas which stand out
because of their ecological, scientific, educational, economic, and political value, in order to
conserve their resources and furnish today's generations with an improved and sustained benefit
from their use, while at the same time, maintaining their potential to meet the needs and
aspirations of future generations;

The Ministry of Agriculture and Livestock — through the Department General of Forest
Development (the present-day Department of National Forests), and by means of the technical
document entitled "Preliminary Strategy for the Conservation of Outstanding Wild Areas of
Ecuador," which was issued in 1979 in the execution of the Project UNDP/FAO-EC/71/527 — grants
the Galapagos Archipelago the highest priority within the National System of Protected Areas, due
to its large number of unique features, including its terrestrial and aquatic ecosystems;

The High Commission encharged with the Revision of the Master Plan for Social
Development and Conservation in the Galapagos Province — in their observations in the stated
documents, as well as in the Immediate Action Plan for the Galapagos Province, promulgated by
Executive Decree No. 937 of July 1 1th, 1985, and published in the Official Register No. 297
(Supplement) of October 22nd of the same year — recommend that the marine zones of the
Galapagos Archipelago be incorporated into the Natural Areas Patrimony;

The Department of National Forests (the official organization responsible for the
administration of the State's Natural Patrimony Areas) — based on studies performed by national
and international organizations — recommends the establishment of a Protected Marine Area within
the sea that surrounds the Galapagos Archipelago for ecological, economic, scientific, educational,
and political reasons and

By virtue of the power previewed in article 78, literal a) of the Political Constitution and
article 69 of the Forest, Natural Areas Conservation, and Wildlife Law.

Decrees

Art. 1 — That the water column, the seabed, and the marine subsoil of the sea located within
the interior of the Galapagos Archipelago — which is understood to be the area within the baselines
used to measure the territorial sea of the Galapagos Archipelago, according to the Supreme
Decree No. 959-Aof June 28th 1971 as proclaimed in Official Register No. 265 of July 13th
1971 — are declared to be a marine resources reserve, along with a band of 15 nautical miles
surrounding said baselines, and fall under the exclusive domain of the State.

Art. 2 — That for the management and vigilance of the marine reserve, an Interinstitutional
Commission comprised of the following members will be established:

-The Minister of Agriculture and Livestock or his proxy, who will preside over the

Commission;

-The Minister of Foreign Relations or his proxy;
-The Minister of National Defense or his proxy;

-The Minister of Industry, Commerce, Integration, and Fisheries or his proxy;
-The Minister of Energy and Mines or his proxy;
—A representative of the National Development Council, and
—A representative of the National Institute of the Galapagos.

The Commission may request the assistance and collaboration of the Charles Darwin
Research Station and national and international organizations as deemed necessary.

Art. 3 — The Ministry of Finance and Public Credit shall make the budgetary transfers
required to the Ministry of Agriculture and Livestock for the adequate functioning of this
Commission.

Transitionary Provision

Within the space of 360 days — starting from this date — the Interinstitutional Commission to
which Art. 2 of this Decree refers shall work out a Management Plan for the Marine Reserve which
will specify aspects relevant to its administration, management, development, and control.

The Ministers of Foreign Relations; of National Defense; of Agriculture and Livestock; of
Finance and Public Credit; of Industry, Commerce, Integration, and Fisheries; and of Energy and
Mines are encharged with the execution of this decree, from the date of its publication in the
Official Register, at which time its enforcement shall go into effect.

Given in Quito, in the National Palace, on the 29th of April, nineteen hundred and eighty six.
Signers:

Leon Febres Cordero Ribadeneyra, Constitutional President of the Republic; Edgar Teran,
Minister of Foreign Relations; Medardo Salazar Navas, Minister of National Defense; Marcel J.
Laniado, Minister of Agriculture and Livestock; Francisco Swett Morales, Minister of Finance and
Public Credit; Xavier Neira Menendez, Minister of Industry, Commerce, Integration and Fisheries;
Javier Espinosa Teran, Minister of Energy and Mines.

An authentic copy — certified and signed by:

Joffre Torbay Dassum, Attorney, Secretary General for Public Administration

(Final translation/R. Lester/June 11, 1986)

A Promise to the Sea,

and the Politics of the Decree

by Roque Sevilla

^achel Carson, in her book The Sea Around Us,
explains human attraction to the sea as stemming
from the fact that our blood has exactly the same
mineral and water composition as the sea.

Whatever the cause, many people have a
great affinity with the sea. They are cheered by its
color, soothed by its murmurs, and awed by its
immensity. I personally belong in this category. I
have always felt very much at ease by the sea,
whether playing in the waves or taking long strolls
down a solitary Ecuadorian beach, watching the
spectacular sunsets so unique to countries facing the
ocean.

My feelings toward the sea became even
stronger the day I descended for the first time with
my SCUBA equipment into deep water. Suddenly I
felt immersed in something to which I was
profoundly attracted. At that moment, I was
absolutely united with my surroundings, a part of the
whole, a sensation I had never felt before.

The Palacio de Corondelet in Quito, center of most important
Ecuadorian political decisions. (Photo by Ignacio de Quadras,
Quito)

Although I was born and raised in Quito,
Ecuador, a city which is cradled in the majestic
Andes, 300 kilometers from the coast and 2,850
meters (9,350 feet) above sea level, I was able
nonetheless to cultivate this sensation. I have since
gone to much trouble to repay the sea for some of
the pleasure it has given me.

A Promise

Two years ago, while strolling down an abandoned
beach, I felt extremely depressed, confronted with
the presence of so much garbage and waste. I
formally promised the sea that I would do anything
possible to protect it from such abuse.

A few months later, while cruising the
Galapagos Islands on one of my diving trips, I met
another lover of the sea, a peaceful and courteous
Englishman who has dedicated most of his life to
revealing the submarine wonders of the Galapagos
to the world. Whenever Godfrey Merlen (see article
page 20) had an opportunity, he would "lobby" for
the sea, trying to convince authorities to intervene in
behalf of its protection.

It was from Godfrey that I heard for the first
time that Gerard Wellington, an American marine
biologist from the University of Houston, Galveston,
Texas, had written in 1974 an extensive study on the
submarine life of the Galapagos. The conclusions of
Wellington's study were fascinating: he observed an
incredibly high rate of endemism (species found only
in a particular locality or region) among the aquatic
species, 35 percent for algae and seashells, and 25
percent for fish. Apparently, marine organisms in the
Galapagos have been forced to adapt to exceptional
conditions because of their relative isolation in the
sea, resulting in both a great diversity of species and
the endemism. As a conservationist and a diver, I
was particularly struck by the percentages, and was
then totally motivated to explore such a fascinating
topic. Back in Quito, I tried hopelessly to find
Wellington's study, The Galapagos Coastal Marine
Environments. As usual, this type of material was not
available to the general public.

National Forestry Director

A strange coincidence gave me direct access to the
source of this report, as I was named National
Forestry Director of the Ministry of Agriculture. The
National Parks Division fell directly under my office.
One of my first satisfactions was to read Wellington's
study, which had fallen into total obscurity among
piles of bureaucratic documents.

The report stated that the protection of the
Galapagos Islands themselves is not enough, and

The Galapagos: a high rate of endemism. (Photo by C. M. Wellington/© National Geographic)

that the archipelago must be considered as an
integral ecosystem with its land and sea components.
Since many animals depend directly or indirectly on

marine resources for their subsistence, it is
absolutely necessary to protect the seas around the
islands as well as the islands themselves.

The National Parks Department and the
Charles Darwin Research Station had been for some
time asserting the need to extend the Galapagos
National Park into the marine area. These demands
were not heard — or more plausibly, there was no
mediator to translate scientific and conservationist
arguments into a language comprehensible to
politicians and authorities.

Mediator

I have spent more than 20 years selling non-tangible
products and services — I started off as an agent in
the stock market, then became an insurance broker.
When I realized how important it was to "sell" the
idea of protecting the Galapagos marine area to key
decisionmakers, I knew the time had come for me
use my skill as a mediator to pay my tribute to the
sea.

At that time, the Woods Hole Oceanographic
Institution (WHOI) had undertaken a study on
coastal and marine resource management in
Ecuador. By invitation, WHOI had decided to
organize an international workshop in the country,
with the participation of highly regarded specialists,
to elicit support for the creation of a marine reserve.

But before the implementation of this plan, as
Forestry Director, I lobbied the Minister of
Agriculture, Marcel Laniado. The Minister was
extremely open to the initiative and offered us his
total support. We then immediately convened with
the scientific community to elaborate on the
technical justifications, background information, and
objectives of the proposed marine reserve area. In
the meantime, I contacted the President's legal
advisor, a lawyer very supportive of conservationist
issues, with whom we had worked on a preliminary
version of the Presidential decree.

Success

After various meetings with government officials and
presidential advisors, the final draft was approved.
On April 29, 1986, the President of the Republic and
six of his Ministers of State signed Decree 1810,
declaring "the column of water, the seabed, and
marine subsoils located within the Galapagos
Archipelago, plus a 15-nautical-mile zone
surrounding the Islands" to be a marine resources
reserve (see decree page 4).

This event received national media coverage
and was applauded by all sectors of the country.
Prince Philip of England, President of the World
Wildlife Federation (WWF) congratulated the
President of Ecuador on this initiative and assured
him that "the WWF considers it a high priority to
protect and manage the extraordinary environment
of the Galapagos Islands."

At present, firm steps have been taken to
implement programs and projects in the marine area.
The Marine Management Plan is being drawn up,
and WHOI, the U.S. National Oceanic and
Atmospheric Administration, the University of Rhode
Island, and The Great Barrier Reef Marine Park
Authority of Australia have offered their technical
support.

A Difficult Road Ahead

This does not mean, however, that we can be sure
that the marine environment of such an
extraordinary region will be sufficiently protected.
There is still a long way to go and enormous
obstacles to overcome. The first obstacle is financial.
Ecuador is a small Third World country with urgent
problems that must be solved immediately. Very
often it is difficult to convince decisionmakers of the
need to invest in long-term conservation measures
when so many urgent human problems demand
immediate responses. In the last four years, my
country has suffered tremendous social and
economic damage from the 1983 and 1987 "El Nino"
phenomena (see article page 42). With the present
fall of oil prices, our national income has been
seriously reduced. Recently we have suffered the
devastation of a major earthquake that killed many
people and destroyed more than 50 kilometers of
our oil pipeline, interrupting our present oil exports
for almost a semester. Nevertheless, I am personally
convinced that Ecuador will try to meet its
responsibilities in protecting the Galapagos Islands
marine area. But, today, more than ever, it will need
international support to fulfill its goals.

Another major obstacle is the Ecuadorian legal
structure. My country has a great diversity of laws to
respond to various needs, interests, and sectors.
According to different circumstances, regulations are
passed by Congress (laws), by the President
(executive decrees), by the Secretaries of State
(ministerial decrees), or by the Municipalities (bills).
Many of these laws, decrees, and bills overlap and
contradict themselves or hide loopholes that have
been overlooked. The regulations that govern the
Galapagos National Park and the urban areas of the
islands are an example of the complexity of the
Ecuadorian legal system. Nongovernmental
organizations in Ecuador will have to play a very
active role to overcome these legal obstacles.

The work of such nongovernmental
organizations is above political and circumstantial
pressures, and many times has proven to be more
effective than governmental activities. My efforts in
protecting the ocean and the Galapagos Islands
would not have been possible if it had not been for
the active support of Fundacion Natura, Ecuador's
most important nongovernmental conservation
organization. The presence of such organizations in
the Third World is becoming ever more prominent;
they have developed great skills in promoting
change and many times have proven to be more
efficient, effective, and flexible in dealing with
problems than governments have. The main reason
for this is that the men and women who create,
sponsor, and work in nongovernmental organizations
are highly motivated and believe in their work. The
protection of the seas and the conservation of the
biosphere will be possible in the long run only
through reliance on local communities and on these
type of organizations.

Roque Sevilla is an economist, a member of the World
Wildlife Federation International Council, and President of
Fundacion Natura in Ecuador.

The Galapagos Marine

Resources Reserve
and Tourism Development

by James M. Broadus

e of the newest and largest additions to the
world's growing treasury of marine reserves, the
50,000-square-kilometer* Galapagos Marine
Resources Reserve, may also be the most fascinating.
The virtually pristine Galapagos marine environment
is remarkable for the abundance and variety of
wildlife inhabiting it, as well as for its unusual mixture
of prevailing oceanographic conditions. These
features have been described by G. M. Wellington
(1984), who highlighted the high proportion of
unique species, the diversity of habitats and species,
the odd assortment of biogeographic kinships, and
the scientific importance of this oceanic area.

The new reserve is fascinating, however, not
only because of the spectacular nature of the
Galapagos marine environment, but also because of
the management issues at stake and the process
through which they are being addressed. The basic
issue is how to assure protection of this special
place, while recognizing the demand by tourists for
access to its wonders, and the desires of local
inhabitants to benefit economically. Declaration of
the Galapagos Marine Resources Reserve grew out
of a master planning effort for the entire archipelago,
and the difficult task of devising a practical
management plan for the reserve provides an
opportunity to improve the integration and
effectiveness of tourism regulation and development
efforts with fundamental conservation principles.

Artificial pressures on the Galapagos marine
environment now appear relatively minor. They
include some danger of small oil spills or cargo loss,
trash from tourist vessels, runoff from coastal
development, collecting of black coral, periodic
operations of a commercial tuna fleet, and artisanal
fisheries — primarily for bacalao (Spanish name for a
type of grouper) and lobster. In the near term, at
least, it is hard to imagine human activities imposing

* The internal waters of the archipelago amount to approxi-
mately 50,000 square kilometers (see map page 2). The
external buffer zone adds some 20,000 square kilometers for
a total area of approximately 70,000 square kilometers.

stresses on the marine system nearly as severe and
disruptive as the natural perturbations of such large-
scale events as El Nino (see article on page 42).
However, the Galapagos biota has long coped with
such natural variations, while human influences are
more likely to be of an entirely new kind. Over a
span of decades, it is likely that uncontrolled human
activities would lead to unacceptable or irreversible
effects. The presently undisturbed quality of the
marine area is therefore a major rationale for
creation of the reserve. This is a chance to
perpetuate a significant unspoiled marine area
before it has been subjected to heavy stress and
alteration.

The Success of Marine Reserves

Establishment of marine parks and reserves has
become a popular means of achieving a number of
national and international goals for marine areas. The
typical rationales for marine parks or reserves have
been described by G. C. Ray (1976) and include
protection of vital habitats; species preservation or
conservation of genetic resources; provisions of
research areas and assurance of comparative
baseline data; recreation, education, and aesthetic
goals; historical or cultural purposes; and an array of
multiple-use resolutions. As early as 1962, the World
Conference on National Parks invited "governments
of all those countries having marine frontiers, and
other appropriate agencies, to examine as a matter
of urgency the possibility of creating marine parks or
reserves to defend underwater areas." To date,
nearly 1,000 coastal and marine protected areas
have been established or seriously proposed
worldwide, although many fewer than this have
been fully implemented.

Habitat protection almost always plays a
major role in marine reserve design; and breeding,
nursery, and refuge zones for resident species are a
common element. Scientific and educational values
often receive greater weight than commercial
economic values, although great emphasis is
typically placed on maintaining recreational
opportunities. The aesthetic appreciation enjoyed by

Workshop on the Role of Science

workshop on scientific research and the
Galapagos Marine Reserve was held from April 20
to 24, 1987, in Guayaquil, the principal port of
Ecuador. The international forum addressed the
role of scientific and technical information in
structuring the reserve, and the needs of the
scientific community in conducting future
research within the reserve.

Nine North Americans and more than 30
Ecuadorians, representing a diversity of scientific
and technical institutions, participated in the
workshop, which was jointly sponsored by the
Marine Policy and Ocean Management Center of
the Woods Hole Oceanographic Institution and
the Oceanographic Institute of the Ecuadorian
Navy (INOCAR).

In opening remarks, Arthur G. Gaines, the
Policy Center's coordinator of foreign planning for
the workshop, stated that "the process of
formulating a workable management plan
ultimately will involve several disciplines: law,
economics, public administration, and others.
Our central objective in the workshop is to
address only one of these — the place and role of
scientific research. From the composition of the
Inter-Institutional Commission and its Technical
Committee [see decree page 4], it is clear that the
importance of science is recognized at the highest
levels of government in Ecuador."

The Director of INOCAR, Lt. Cmdr. H. R.
Moreano, summarized the work of his institute at
a concluding session:

"First, the hydrographic surveys to update
the charts of the Galapagos waters began some
years ago and some new charts are now

available, such as those of Wreck Bay, Academy
Bay, Plaza Islands, Puerto Villamil, etc. Our
research vessel, the R/V Orion is presently
surveying the area of Bolivar Channel, Banks and
Urvina Bay, Punta Espinoza, and Tagus Cove.

"Second, the Oceanographic research,
especially on the west side of the archipelago
(92 degrees W), has collected almost 8 years of
data with the goal of understanding the
upwelling and circulation problems in the area
and its implication in primary and secondary
productivity.

"The last project, started in mid- 1 986,
includes a station just south of Academy Bay for
recording biological and physical Oceanographic
data. This project is being coordinated with the
Charles Darwin Research Station.

"But the point I want to emphasize is that
the information INOCAR is getting through these
projects is not enough for a complete
understanding of the Galapagos marine
environment. This is mainly due to a lack of
equipment — which is one area where
international cooperation could come in. For
example, during hydrographic surveys, the R/V
Orion could collect seismic, magnetic, and
gravity data. This would be useful for a better
understanding of the geological setting of the
Galapagos Islands. Surface and subsurface current
measurements using a profiler current meter,
buoys, etc., could help to improve our
understanding of physical problems.

"If any foreign scientists or institutions are
interested in carrying out research in Galapagos
waters, a starting place would be to write a letter

visitors is also frequently a priority. Marine reserves
are more likely than onshore reserves to be designed
to accommodate multiple uses, and the assurance of
continued traditional and artisanal uses of marine
areas is a characteristic of many marine reserves.
Whatever the relative weight placed on specific
objectives, establishing a marine reserve can provide
an organizational and administrative framework for
the rational balancing of uses and goals in areas
where such structure is otherwise absent. A
common standard of success in many reserves is,
conservatively, merely to continue the status quo
while protecting marine ecosystems and water
quality.

In most instances, the goals of marine reserve
establishment are quite long-term in nature. Despite
the relative novelty of marine reserves, several
striking examples of success can be identified
already. One of the oldest marine protected areas is
in the United States at Key Largo in Florida. There,

an impressive roster of accomplishments have been
witnessed in terms of habitat protection, reef
recovery from perturbation, recreational safety and
management, commercial opportunities, and
education. In the southern Sinai, establishment of a
marine reserve by the government of Israel
effectively eliminated the destructive fishing
technique of dynamiting reefs. The reserve also
created the basis for a healthy tourism industry in a
remote area, and the government of Egypt is
contemplating expansion of reserves for similar
purposes. In Australia's Great Barrier Reef Marine
Park, economic benefits have accrued from the
combination of carefully regulated tourism with
enhanced protection of natural environmental assets
(see Oceanus Vol. 29, No. 2).

The prospect of similar benefits helped
motivate the May 1986 declaration of the Galapagos
Marine Resources Reserve. The reserve presents an
opportunity, for the first time, to manage the

in the Galapagos Marine Reserve

to the Director of INOCAR expressing the intent
of their research.

Final workshop recommendations were:

1) Science should influence the framework of
the Marine Reserve management plan in two
distinct ways:

a) Scientific information and
methodologies serve as the basis for
defining physical realities of the natural
and cultural systems addressed by
management.

b) Reserve Management should
incorporate the needs of Ecuadorian and
foreign scientists and scientific
organizations for conducting future basic
and applied research.

2) National and international scientific
cooperation should be promoted as a means to
fund and carry out the large number of
research, survey, and monitoring studies needed
for proper management of the reserve. The
great international appeal for funding of science
in the Galapagos archipelago stems from the
union of:

a) Unusual natural features of the setting.

b) Prospects for international collaboration.

c) Ecuador's rededication to conservation
of natural systems for science,
education, and low impact use.

3) Because of the great size and complexity of

the Marine Resources Reserve, remote sensing
techniques should be used fully to define the
environment, its resources and habitats, human
impacts, and El Nino variability. More
specifically:

a) Existing imagery should be obtained and
made available to Ecuadorian scientists.

b) The Administrator of the U.S. NOAA
should be urged to support launching of
a new Coastal Zone Color Scanner
satellite to replace the non-operational
one.

c) Large-scale aerial photographs (color
stereo pairs) of the entire Galapagos
coast are needed for many purposes in
managing the reserve: Ecuadorian
agencies should use existing facilities
and expertise to obtain these photos.

4) Public participation should be encouraged in
setting goals and priorities for management of
the reserve. An enhanced sense of stewardship
should be instilled in all Ecuadorians with regard
to the Galapagos archipelago.

5) The focus of these recommendations-
science — is only one aspect of management
formulation. The government of Ecuador should
support the future efforts of the Inter-
Institutional Commission and its technical and
advisory committees in addressing other
management components for the Marine
Resources Reserve. — PRR

archipelago's ecosystem as a whole, and to fulfill the
objective of its Biosphere Reserve status in the
United Nations' International Man and Biosphere
Program. This program seeks the protection of
ecosystems representative of the biogeographical
regions of the world, while also incorporating human
activities. Authority of the Galapagos National Park
(PNG), established in 1959 and incorporating some
90 percent of the islands' land area, was limited to
onshore areas only. No mechanism existed to
coordinate policies for the highly interdependent
marine and terrestrial systems. Neither was there a
clear framework within which to regulate human
activities in the marine area and to assure their
compatibility with national goals for environmental
conservation in the archipelago.

Genesis of the Galapagos Marine Reserve

Naturalists have long marveled at the wonders within
Galapagos waters (see box page 22), and a marine
component for the Galapagos National Park has

been envisaged since at least 1973. This is reflected
in the 1974 Master Plan for the PNG, which called
for an extension of Park boundaries 1 kilometer
seaward of all uninhabited islands (only four of the
archipelago's 19 islands are inhabited). A similar
proposal was made at the Charles Darwin Research
Station by G. M. Wellington in 1975, though his plan
called for a seaward extension of 2 nautical miles
from all islands and specified in some detail a marine
zoning scheme. Partly in reaction to restrictions
proposed in Wellington's zoning scheme, strong
local opposition arose to resist the marine park
proposal. In 1978, there were reports that the
government of Ecuador intended to extend PNG
boundaries 15 miles from all islands and to include
all internal waters of the archipelago, but no action
followed. Three years later, a High Level
Commission to Study the Impacts of Tourism in
Galapagos recommended a PNG extension of 1 to 5
kilometers around all the islands.

Local opposition to the idea may have been

tempered a bit by then President Osvaldo Hurtado's

1982 speech declaring that an objective of a reserve
would be not only to protect ecosystems, but to
protect the islands' marine resources for the
privileged use of local residents. This was followed in

1983 by a very moderate and permissive zoning
proposal from Darwin Station marine biologist Gary
Robinson that left virtually all traditional uses
unaffected. Robinson suggested a 2-nautical-mile
marine extension of PNG boundaries, but he also
argued that inclusion of all internal waters would be
best.

Researchers from the Woods Hole
Oceanographic Institution (WHOI) also played a role
in creation of the Galapagos Marine Resources
Reserve. In 1983, a team from WHOI's Marine
Policy Center was invited by the government of
Ecuador to examine the status of coastal and marine
resources management in the Galapagos. The WHOI
team's 1984 report to the commission that was
devising a master plan to balance conservation,
tourism, and development, included this finding:
"While national policy for the onshore area of the
Galapagos Islands unambiguously stresses
conservation of the natural environment, no clear
policy seems to have been defined for the
Galapagos marine area. The long-pending issue of a
marine park or reserve for the Galapagos is one of
the major questions facing coastal area and marine
resources management there."

Among several policy options identified by
the WHOI team for consideration by the
government of Ecuador, were the following:

Establish policy that future creation of marine
park or reserve in Galapagos waters is envisaged
and initiate discussions and further study toward
most appropriate definitions and timely
implementation of such a park or reserve.

Establish a marine park or reserve in the
Galapagos immediately by decree, which could
leave open the details of the organization and
implementation of the park or reserve.

As it turned out, this was more or less the
course followed by the government. In July 1985,
President Febres Cordero adopted, as national policy
a "Plan for Immediate Action" that included:
"Preparations should be made for an Executive
Decree incorporating a marine reserve into the
PNG." Further studies were pursued in the
meantime, and in May 1986 the Presidential decree
was issued (see page 4). During the process leading
up to the decree, crucial political leadership was
provided by Roque Sevilla, then director of the
National Forestry Agency and an advisor to their
Minister of Agriculture, Marcel Laniado. Gunter
Lisken, the country's Subsecretary for Industry and
Tourism, also played a prominent role in establishing
the policy, as did individuals in the Navy and Foreign
Ministry. Interestingly, the reserve's 15-mile external
band may have helped generate enthusiasm with
some members of Ecuador's foreign policy
establishment. They saw possible support for

Ecuador's claim to a 200-nautical-mile territorial sea
in the extension of control beyond the 12 nautical
miles normally recognized for territorial seas in
international law.*

The Central Issue — Tourism

Such subtle issues of international law, however
important to Ecuador's national security concerns,
do not approach the immediate practical importance
and contentiousness of the central issue in the
Galapagos: Tourism. Organized tourism has existed
since the mid-1960s. It grew up within a framework
of management and regulation similar to that for the
PNG and the Darwin Station.

Indeed, for a number of years the tourism
industry was itself an important force for
conservation in the islands. The cooperation
between tourism interests and environmentalists was
even cited as an example for other natural areas with
high tourism appeal. In recent years, however,
indigenous pressures for economic development
and associated political events have begun to disrupt
this constructive relationship.

Confined to small "colonized" areas on four
islands, and surrounded by arid national parklands
and the Pacific Ocean, the rapidly growing
Galapagos population is hard pressed for a
livelihood.

Tourism now surpasses traditional farming and
fishing activities as a source of employment. The
annual number of visitors increased from about
4,500 in 1970 to more than 26,000 at present, with
an average annual growth of 24 percent between
1976-1980.

Aside from worries about the direct effect on
the environment from a growing volume of tourists,
tourism growth also has the indirect effect of
attracting to the islands new permanent settlers
seeking a place in the industry. The permanent
population now numbers more than 8,000, and in
recent years the two largest colonized areas, on
Santa Cruz and San Cristobal, have experienced
annual population increases of about 10 percent. A
ceiling on the annual number of visitors was set at
12,000 in 1973, but this was apparently surpassed in
1978. A new ceiling of 25,000 visitors was
established in 1982 and confirmed in the revised
master plan of 1985. This ceiling also has been
surpassed. There is some pressure to raise the ceiling
and little attention to the implications for permanent
population increases and associated needs for
additional sources of local revenue.

Quite obviously, management of the Marine
Reserve will be affected by policies governing tourist
access to the archipelago. By the same token,
regulatory decisions on uses of the marine area will
affect the tourism industry. The tourism experience
in Galapagos has always been waterborne and

* Under international law, most states claim a territorial sea
12 nautical miles in width. In these waters, the coastal state
has complete sovereignty, except for rights such as
innocent passage and certain other historical rights. The
generally common 200-nautical-mile exclusive economic
zone (EEZ) implies coastal state control of the resources of
the water, seabed, and subsoil.

LEGEND

i r I Arrows show
~jjj direction of
plate movement

Middle America Thrust

The Galapagos Islands are the tips of huge submarine volcanoes formed as a crustal plate passes over a mantle "hot spot." /As a
result of sea-floor spreading, the islands are moving south and east at more than 7 centimeters per year. The southeastern island
of Espanola has the oldest dated rocks at 3.25 million years, while to the west, Fernandina and Isabela are less than 0.7 million
years old. (After M.H. lackson, Galapagos: A Natural History Guide, 1985)

water-based. A typical tourist visit involves arrival by
air (though some come by cruise ship) and almost
immediate transfer onto a boat for a week or two-
week tour of the islands. There are now three cruise
ships (50 to 90 passengers each) operating in this
way, and some 45 smaller vessels (6 to 15
passengers). Many are converted fishing boats. In
1971, there were 6 small tour boats, and 1 cruise
ship.

The spectacular worldwide boom in marine
recreation suggests that increasing demands for
access to Galapagos waters will be expressed by
tourists, and an incipient diving services trade is
already emerging (see article page 20). However,
there is still no mechanism for the regulation and
control of such activities, and there is only a limited
capability to assure the safety of diving visitors or to
respond to accidents should they occur. Increasing
tourist use of Galapagos waters also opens issues
about the growth and control of more traditional
water sports, such as spear and sport-fishing,
sunbathing, recreational boating (including
windsurfing), and waterskiing.

The temptation for Ecuador to relax

In the Galapagos, a stationary hot spot in the mantle gives
rise to volcanoes as the sea floor moves over it. Cycles of
volcanic activity leave a trail of volcanic peaks, which
become older and more eroded in the direction of plate
movement. (Source: M.H. lackson, 1985)

Cruise ships bring large numbers of tourists into the new Galapagos Marine Resources Reserve, and serve as floating hotels
during their stay. In addition, smaller boats can be chartered locally for day and weekly trips throughout the islands. (Photo by
Arthur Gaines)

restrictions on tourism in the Galapagos must be
great, for foreign revenues are at a premium and
tourism is a major source. In 1980, for example,
when earnings from petroleum and shrimp exports
were high, tourism still generated 22 percent of the
country's foreign earnings, and almost a fifth of that
was from Galapagos tourism. Depending on the
nature of demand for Galapagos tourism, however,
some economic principles suggest that greater
revenues can be earned with more restricted access
and tighter controls on the market. It also has been
suggested that Ecuador has many mainland tourist
attractions that might be developed to rival the
Galapagos, thereby taking some of the tourist
pressure off the islands.

During the early years of tourism in the
islands, the industry was dominated by one Quito
company. In recent years, there has been a huge
amount of new tourism activity. The small boat
sector has grown rapidly and the variety of package
tours available is proliferating. No longer must
tourists "take it or leave it" with expensive, week-
long boat tours. So called "economy tourism" is
growing, based on hotel accommodation and day
trips to nearby islands. The tourists gain, of course, in
terms of savings and flexibility. The uncontrolled
result for operators and the Galapagos, however,
may be a maximum number of tourists and virtually
no tourism profit. With associated growth in

population, there must be a real concern that this
process will eventually spoil the natural attraction
that draws tourists in the first place.

Prognosis

Two years ago, I wrote in these pages (Oceanus Vol.
28, No. 1, Spring 1985) that "no proposal for the
Galapagos marine area appears to be moving
forward." Even then, however, the situation was
hopeful: "The ingredients for progress in devising
appropriate management provisions for the
Galapagos marine area seem to be in place. This
pristine area is of great scientific interest and of vital
importance to the archipelago's terrestrial organisms.
A scientific research station is already at the scene
and functioning, and the station's marine research
capabilities have been upgraded. A new national
government is turning its attention to the situation,
which commands broad public interest. A national
park is a major presence onshore, and the Navy
already maintains an administrative and enforcement
structure in the archipelago. The Galapagos Islands
themselves enjoy high international visibility. If a
comprehensive management program cannot be
devised and successfully implemented for this
watery treasure, we must surely be pessimistic about
what can be achieved for other such marine areas
elsewhere in the world."

Great progress has been made in the

About 26,000 tourists visit the Galapagos yearly. Some developers seek an increase to 150,000 — a number that would likely
seriously stress the islands' ecosystems. (Photo by Andrew Rakoczy, National Audubon Society, Photo Researchers)

intervening two years. With the declaration of the
Galapagos Marine Resources Reserve, an optimistic
step has been taken. Details of the reserve's
implementation must still be devised, but the policy
and direction are now clear. Realistically, the goals
and the planning horizon are long term. As an
example, a management plan for the PNG, which
has done so much to protect the islands, was first
completed in 1974. The PNG itself, however, was
originally established in the mid-1 930s. For the
Galapagos marine area, the burden of proof has now
been shifted in favor of comprehensive management
and protection. What this means ultimately in the
face of tourism and development pressures waits to
be seen.

lames M. Broadus is Director of the Marine Policy and Ocean
Management Center at the Woods Hole Oceanographic
Institution.

Letter Writers

The editor welcomes letters that comment on
articles in this issue or that discuss other mat-
ters of importance to the marine community.

Early responses to articles have the best
chance of being published. Please be concise
and have your letter double-spaced for easier
reading and editing.

Selected References

Beebe, W. 1924. Galapagos: World's End. New York and London: C.

P. Putnam & Sons.
Broadus, ]., I. Pires, A. Gaines, C. Bailey, R. Knecht, and B. Cicin-

Sain. 1984. Coastal and marine resources management (or the

Galapagos Islands. Woods Hole Oceanographic Institution Tech.

Rept. WHOI-84-43.
Broadus, J. 1985. Poor fish of Redondo!: Managing the Galapagos

waters. Oceanus 28(1) 95-99.
Budowski, G. 1976. Tourism and conservation: Conflict, coexistence

or symbiosis? Environmental Conservation 3(Spring): 27-31.
Clark, E. 1977. Synogogues and sea fans: Israel's national parks and

nature reserves. National Parks and Conservation 51: 13-20.
Davis, G. 1981. On the role of underwater parks and sanctuaries in

the management of coastal resources in the southeastern United

States. Environmental Conservation 8: 67-70.
Garces, F., and ). Ortiz. 1983. El Turismo en El Ecuador y su Relacion

con Galapagos. Quito, Ecuador: INGALA.
Ray, G. C. 1976. Critical marine habitats: definition, description,

criteria and guidelines for identification and management. Proc.

International Conference on Marine Parks and Reserves, Tokyo,

Japan. IUCN Publications New Series 37.
Robinson, G. 1983. A Marine Park in the Galapagos. Noticias de

Galapagos 37: 9- 13.
Silva, M., E. Gately, and I. Desilvestre. 1986. A bibliographical listing

of coastal and marine protected areas: A global survey. Woods

Hole Oceanographic Institution Technical Rept. WHOI-86-1.
Wellington, G. 1984. "Marine Environment and Protection." In, Key

Environment Series: Galapagos Islands, ]. E. Treherne and R.

Perry, eds., pp. 247-264, Oxford: Pergamon.

Two Legal Opinions on the

Ecuadorian Law

by Efrain Perez Camacho

In Ecuador, in the past century, there was already a
consensus on the need of a special status for the
Galapagos Islands. The Ecuadorian Constitution of
1883 calls for special laws for the Colon
Archipelago — as the Galapagos used to be called in
legal documents.

The most visible Ecuadorian politician of this
century, J. M. Velasco Ibarra, four times president,
offered to design a coherent set of rules for the
management of the archipelago in 1955. "Such a law
will include every aspect, among others economics,
social, administrative ones, etc., to solve the multiple
problems of the insular territories." Alas, such a
purpose was not, and has not been fulfilled.

On the other hand, by bits and pieces, several
statutes have been introduced expressly for the
Galapagos on different subjects ranging from
protecting and managing the park to public
administration and public servants' salaries. And
because the islands are a part of the Ecuadorian
territory, most of continental Ecuador laws and rules,
many of which encourage development, do apply.
So, for the Galapagos, we encounter a dual set of
rules: those special for the islands, and those general
for the country.

On May 13, 1986, the Ecuadorian
Government established a reserve for marine
resources in the Galapagos. It included the
archipelago's interior sea, a 15-nautical-mile
surrounding buffer zone, the water column, and the
seabed and its subsoil. A committee was established
to oversee and control the reserve; its members are
representatives of the following state secretaries:
Agriculture (park services), Foreign Relations,
Defense, Industries, and Fisheries (oversees tourism
and fisheries), Energy and Mines, Planning, and the
National Galapagos Institute (INGALA).

The Decree mandates that a management
plan be made, which should address policy,
management, development, and control of the
marine reserve. The Forestry Law (decree 1529, 22
February 1983) calls for specific management plans
for State natural areas that should contain, among
others, the following items: basic information,
inventory of the area, verification of boundaries,
objectives of the area, zoning, programs of

interpretation and research, direction, management,
and protection of the environment.

The task of the management plan for the
marine reserve in the Galapagos is a challenging one,
both because of the great area involved and because
it will be the first of its genre. The management plan
for the Machalilla terrestrial and marine park, on the
coast of continental Ecuador, has been just finished,
but it can not be said that its circumstances and
ecological environment are similar to the Galapagos,
so the Machalilla management plan will not be of
much use as a background for the Galapagos case.

The Galapagos marine reserve fails under the
provisions of the Forestry Law, Natural Areas and
Wildlife (Law No. 74, 24 August 1981). It is basically
a biological reserve, which is defined in the law as an
area of variable surface, that could be in either the
terrestrial or the aquatic environment and that is
dedicated to the preservation of its species.

Additionally, because of the potential for
mineral resources on the Carnegie Ridge, near the
archipelago, the seabed and the marine subsoil was
included as well into the reserve; so the name is not
just "biological reserve," but rather marine resources
reserve, that encompasses both the living resources
and mineral ones.

In the case of mineral resources, the idea is to
spare the archipelago of the possibly disastrous
ecological consequences of a future industrial
exploitation in great scale.

The Forestry Law does not include among the
categories of natural areas of the State the reserves
of marine resources. But as the establishment of
such a reserve is an administrative matter, it is well
within the authority of the Executive Branch to
create one. What it means is that more flexibility is
given to the concept of reserve; enough for
additional resources to be included in it. So, my
interpretation is that for the Galapagos marine
resources reserve, the Forestry Law applies, plus
additional provisions to be established in the future
through executive decrees.

The presidential Decree of 1986, after
declaring the reserve, charged an ad hoc committee
to come up with a management plan within a six

continued on page 18

Galapagos Marine Reserve

International Issues

by Kilaparti Ramakrishna

On May 13, 1986, the President of the Republic of
Ecuador decreed the "archipelagic waters" of the
Galapagos Islands along with a surrounding band of
waters 15 nautical miles in breadth to be a "Marine
Resources Reserve" falling under the "exclusive
domain" of Ecuador. (See decree page 4).

The decree recounts several steps taken by
Ecuador in the past that accorded highest priority to
the protection and preservation of a large number of
unique ecological features of the islands. A
consensus existed in Ecuador on the need for a
special status for the Galapagos Islands as early as
the 19th century. In fact, the Ecuadorian Constitution
of 1883 called for special laws for the Colon
Archipelago, as the Galapagos were then called in
legal documents.

The 1986 decree, as drafted, raises interesting
international legal issues. These are issues that arise
when any coastal State proposes to designate a given
area as a marine park or sanctuary. The situation is a
little more complex when the coastal State is a
developing country, and the area covered is not
entirely within its internationally recognized
territorial waters.

In this connection, it is important to note that
Ecuador has claimed territorial waters to a distance of
200 nautical miles from the appropriate baselines
both along the mainland and around the Galapagos
Islands and that Ecuador has not signed the United
Nations Convention on the Law of the Sea of 1982
(UNCLOS), which states that the breadth of the
territorial sea, over which the coastal state has
sovereignty, shall not exceed 12 nautical miles.

Therefore, when the Marine Resources
Reserve Decree established Ecuadorian
"sovereignty" over a band of 15 nautical miles
extending seaward from the archipelagic baselines
drawn earlier, it had the effect of extending the
Ecuadorian jurisdiction by 3 nautical miles beyond
the internationally recognized limit for the territorial
sea.

If, as several commentators have pointed out,
the 200-nautical-mile territorial sea in Ecuador and
some other South American states is comparable
with the now recognized concept of "exclusive
economic zone" (EEZ), the 15-mile band would gain

considerable importance. The question that then
needs to be addressed is: What kind of restrictions
does Ecuador plan to impose in these waters? At this
point in time, however, this is unclear.

Assuming that the protection and preservation
of the rare and fragile marine ecosystem of the
Galapagos requires that there be restrictions on
freedom of navigation and on fishing, what
procedures does international law provide to
accomplish this? Does one look to either
conventional law or state practice? The safest way
may be to look at both.

UNCLOS contains a general provision that
States shall take measures necessary to protect and
preserve rare or fragile ecosystems as well as the
habitat of depleted, threatened, or endangered
species and other forms of marine life (article 194.5).
Likewise, if special measures are required for
"recognized technical reasons," the coastal state may
adopt the view that, despite the existence of
international pollution standards, certain areas of
their EEZs display characteristics that call for specific
measures of protection (article 21 1.6.a). This
provision generally is referred to as relating to
pollution from vessels. Read in conjunction with the
extent of the sovereign rights of coastal States to
conserve and manage natural resources in their EEZ
(article 56.1. a), this provision leads some to believe
that UNCLOS provides for the establishment of
special protected areas.

The key determinant, however, is the phrase
"recognized technical reasons." Some government
spokesmen in Ecuador have been reported
explaining that the 15-nautical-mile boundary is
determined by the foraging range of certain
protected species of marine birds based in the
reserve. It is not clear, however, if this is a sufficiently
recognized technical reason. This determination can
be made only by an appropriate/competent
international organization, and the coastal state shall
implement international rules and navigational
practices so recommended by the organization.

In addition to UNCLOS, the other reference
to the establishment of special areas can be found in
the Convention for the Prevention of Pollution from

continued on page 19

months period that could well be extended far
beyond. A management plan has several elements,
like zoning, uses, etc., that we are not covering in
this article, even if they do indeed present legal and
institutional problems. We believe that any short-
term legal and institutional study should immediately
address the legal status of the fisheries and the
restructuring of the public administration in the
islands. As for national fisheries, there are both
artisanal and commercial ones in the Galapagos.

The artisanal fisheries do not pose any
immediate or significant problem as they are now.
The studies conducted on these fisheries seem to
agree that there is not any danger to the species of
the islands that could be caused by such activities. A
thorny issue, that is as ecological as it is economical
and political, will be the very likely situation that
must arise with the improvement of the gear and
reach of those fisheries. Should they still be allowed
to work their trade or what kind of legally
enforceable limitations could be imposed on them?

The Ecuadorian commercial fisheries can not
operate in the area of the reserve. How this will
affect the national tuna fleet and how this
prohibition should be interpreted in its relationship
to the Fisheries Law is not known.

The main institutional problem that the
Galapagos marine reserve poses is an administrative
one. Administrative problems already exist for the
National Park and they will be compounded by the
establishment of the reserve. We have a few
questions that we believe should be urgently
addressed by the management plan:

• The Galapagos National Park has functioned
successfully for most of the archipelago's
terrestrial area for a long time and its
achievements have obtained international praise.
Its main problems are financial and lack of
adequate staff. Good coordination with other
public offices in the islands has not always been
possible. So an immediate question comes to
mind: Should the marine area of the Galapagos
reserve be managed by the same office that is
managing the land area of the National Park?

• We have mentioned the development-oriented
bias of the Municipal Law in Ecuador that is
applied to the Galapagos as well. How can we
adapt the Municipal Law in a way that the
municipalities' actions in the islands intermingle
more appropriately with the Galapagos unique
environment?

• The public offices in the islands have different
objectives. Their actions sometimes contradict
and supersede, creating conflicts and hindering
the work of the park and, assuredly in the future,
the work of the reserve. Would it be more
convenient to join the whole of the state activities
under a single ad hoc authority? In that case,
what should be the nature of the civil servants'
work and salary regime?

• Furthermore, the Forestry Law, Natural Areas
and Wildlife regulation allows for conservation,
education, and research activities in the resources
of the biological reserve. What kind of
arrangement can be established that will allow
scientific research to be translated into better
policies for management of the reserve?

The Ecuadorian legal system does not have an
answer for these questions. Effectively, it has a
developing country juridical structure, that is, one
that encourages and fosters growth and
development. As the Constitution of 1883 put it
plainly more than 100 years ago: We need a
different set of laws and regulations for the
Galapagos. The new Galapagos statute must contain
rules for the adequate improvement of the living
standards of the islanders, delicately balanced with
the preservation of the archipelago's environment:
We should be seeking a true eco-development.

Efrain Perez Comacho, an attorney, is an authority on the
Public Law of Ecuador and has been a Guest Investigator at
the Woods Hole Oceanographic Institution's Marine Policy
and Ocean Management Center.

The new Galapagos Marine
Resources Reserve includes a
15-nautical-mile external
marine buffer to protect the
foraging range of seabirds, such
as these masked boobies.
(Photo by D. /. H. Phillips)

Ships of 1973 (MARPOL Convention). Annex 1
defines "special area" as a sea area where
recognized technical reasons require the adoption of
special mandatory controls over oil pollution. Even
here, however, the determining factors are the
oceanographic and ecological requirements as well
as traffic of a particular character. Accordingly, it may
be concluded that there are difficulties in using this
provision to impose a blanket ban to protect an area.

Other conventions of importance are the
Protocol Concerning Mediterranean Specially
Protected Areas and the Protocol Concerning
Protected Areas and Wild Flora and Fauna in the East
African Region. The first protocol was established in
1982 under the Convention for the Protection of the
Mediterranean Sea Against Pollution (1976). The
second was promulgated under the Convention for
the Protection, Management, and Development of
the Marine and Coastal Environment of the Eastern
African Region (1985). Both protocols make special
provision with respect to the establishment of a
protected area "contiguous to the frontier or to the
limits of the zone of national jurisdiction of another
party," thus requiring consultations between parties
and facilitating international co-operation.

In addition to provisions in these multilateral
conventions, several national legislations also were in
place that relate to specially protected areas in the
sea. Leading examples may be found in the laws of
developed countries such as Australia, New Zealand,
the United States, Britain, and West Germany and in
the laws of some developing countries such as
Antigua and Barbuda, Barbados, and Bermuda. More
detailed study is needed to substantiate whether
available conventional law and state practice bring
out a clearly recognizable general principle of
international law and, if so, what its connotations
may be.

All this notwithstanding, it is reasonable to say
that a substantial body of conventional law and the
makings of customary international law (not a well-
developed body of law at this time) exists for the
protection and preservation of unique and fragile
marine ecosystems. In light of the above, the
declaration by Ecuador establishing a 15-mile band
around the Galapagos is not in itself a major
departure from either the customary or conventional
law. It must, however, be said that Ecuador has as
yet done very little to gain acceptance from the
competent international organizations for any
"recognized technical reasons."

Treating the terrestrial and marine
environments as one large marine ecosystem and
adopting an ecosystem concept of management in
Galapagos is urgently needed. At the same time, the
Government of Ecuador should consult with the
concerned/competent international organizations in
obtaining wider support of the Galapagos as a
specially vulnerable area and convincing them that
special regulations are warranted to safeguard its
environment for future generations. In the present
case, the concerned/competent international
organizations include the International Maritime
Organization, and the Food and Agriculture
Organization. It is true that these organizations may
take anywhere between two to three years to bring
into effect any of the accepted special regulations.
Ecuador in the meanwhile would have some valid
legal bases to impose the required restrictions.

Kilaparti Ramakrishna is a Fellow at the Marine Policy and
Ocean Management Center, Woods Hole Oceanographic
Institution. He is also a Visiting Scholar at the Harvard Law
School, Cambridge, Massachusetts, and is on leave from the
Indian Society of International Law, New Delhi, India.

Author's rendition of modern lobster fishing in the
Galapagos, using surface-supplied air. This method consists of
the use of a normal regulator supplied with air from a
compressor carried on a small dinghy, which is equipped
with an outboard motor.

Diving in the Galapagos

By Godfrey Merlen

I he 19th and 20th centuries have opened the door
to exploration and exploitation of the marine
environment. Until then, investigations beneath the
water had been limited to a few adventurous
humans using primitive equipment and risking life
and limb in the process. The Industrial Revolution
made available new materials and machinery
capable of compressing air to compensate for the
tremendous increase of pressure with depth. This
eventually led to the invention of SCUBA (self-
contained underwater breathing apparatus), giving
amateur and professional diver alike the flexibility of
a fish. With the use of a diving mask to restore his
vision to its excellent aerial quality, a new dimension
was opened in which man could exercise his natural
curiosity and manual dexterity.

The extent to which this has occurred during
the last 40 years has led to concern — in certain areas
there is grave danger of divers despoiling marine

environments by over-exploiting marine resources
and physically damaging areas of great beauty.

The Galapagos are now in a critical period
with regard to the direction that man will follow in
his use of the marine environment. In May of 1986
the internal waters of the archipelago (that is, the
waters contained within the figure formed by
connecting the extreme points of land) plus 15
nautical miles to seaward were proclaimed a Marine
Resources Reserve by the Ecuadorian Government
(see map page 2). This is a major step forward, but it
is extremely important that when the laws regarding
the area are established, much consideration be
given to the place of diving for gain and for pleasure.

Laws governing the terrestrial islands can
provide a model. Public access is allowed to the
National Park (established in 1959), but that access is
limited to specific areas and under strict rules to
provide protection for the fauna and flora of these

remarkable islands. The marine subsurface
dimension is as varied and magnificent as the
terrestrial section of the islands, and will require the
same type of planning if the overall integrity of the
island ecosystems is to remain intact. The two
sections are inextricably linked through food webs of
many organisms, ranging from crabs to sea lions.

Earliest Divers

Although there can be no doubt that people were
exploiting the sea for decades or even centuries
before the present, the first instance of diving in the
Galapagos that I can find comes from William
Beebe's The Arcturus Adventure, which deals with his
voyage in 1925 as a part of the New York Zoological
Society's Oceanographic Expedition (see box page
22). This exploratory effort was made with a hard hat

At left, diver in Ca/apagos with porcupine fish (Photo by
T. M. Rioux, WHOI)

'Before One
Could Remember
To Be An
Ichthyologist'

EDITOR'S NOTE:

Charles William Beebe (1877-1962), an
American biologist and natural historian, was a
rare combination of scientist, explorer, and
literary talent. Director of tropical research for
the New York Zoological Society, he led many
scientific expeditions — and among them several
trips to the Galapagos. One of his many books,
The Arcturus Adventure, chronicles his 1925
trip to the islands. In the chapter, "With Helmet
and Hose," he expressed, with typical
imagination and wit, the wonder of his first
diving experiences. (In 1934, Beebe descended
in a bathysphere to a record depth of 923
meters in waters off Bermuda.) A sample of
"With Helmet and Hose" follows:

During the first part of the Arcturus adventure,
the sea was too rough to think of using it, even a
few feet below the gangway, but when we
moored close under the cliffs of Darwin Bay at
Tower Island I brought up the box from the hold
and unlimbered the diving apparatus. The helmet
was a big, conical affair of copper, made to rest
on the shoulders, with a hose connection on the
right side and two oblique windows in front.
Around the bottom extended a flange on which
four flattened pieces of lead were hung, each
weighing 10 pounds. This made a total weight of
60 pounds for the entire thing. The hose, which
was of the ordinary common or garden variety,
was attached at one end to the helmet and at the
other to a double-action automobile pump,
which screwed to a board, and was operated by
a long iron lever, pushed back and forth. . . .

Our regular mode of diving is as follows:
We start out from the Arcturus in a flat-bottomed
boat which has a square, 18-inch glass set in the
bottom amidships. To the stem is fastened a long,
metal jacob's-ladder, rolled up when not in use.
We are towed or we row to the shore, preferably

Beebe in diving helmet.

to the base of cliffs or steep rocks, as that affords
considerable depth close inshore and rocky
places are beloved by hosts of fish. We anchor as
close to the cliffs as is safe, and roll out the
ladder, so that it sways in midwater or rests upon
the bottom. The pump is in the bow, the handle
fixed, and the leather washer carefully screwed in.
The hose is cleared of kinks, and is looped, partly
overboard. A hand line is tied to the top of the
helmet, and the inside of the glass windows is
coated with a film of glycerine to prevent the
breath of the diver from condensing and so
clouding it. The four lead weights are slipped
over the flange on the helmet base and all is
ready for the diver. A hand water-glass is near for
constant lookout for danger, and one or two
long-handled harpoons.

In bathing suit I climb down the ladder
over the stern, and dip to my neck, being careful
not to wet my head. Then lohn lifts the helmet; I
give a last, quick look around, draw a deep
breath, duck into it, and as it settles firmly on my
shoulders, I climb slowly down. The sensation
just above water is of unbearable weight, but the

weighing about 60 pounds, utilizing a hand-operated
compressor located in a small boat. Beebe's
descriptions of the sharks and brightly colored reef
fish are interesting, but one senses a nervousness of
being inside such a cumbersome piece of

equipment, with its extremely limited view of the
outside world.

Commercial Diving for Lobsters

Commercial diving in the Galapagos seems to have

instant I immerse this goes and the weight of the
helmet with all the lead is only a gentle pressure,
sufficient to give perfect stability. . . .

from a blurred view of the water surface
and the boat's stem, I sink instantly to clear vision
under water. I descend three rungs and reach up
for the short harpoon or grains which is put into
my hand. At the fourth or fifth rung the air presses
perceptibly on my ears and I relieve it by
swallowing. This ceases as soon as the helmet is
entirely under water. I descend slowly,
swallowing now and then, and when the last
rung has been reached, I lower myself easily by
one arm, and lightly rest on the bottom. If serious
danger threatens or the pumping should go
wrong for any reason, I have only to lift up the
helmet, duck out from under it and swim to the
surface. The level of the water keeps constantly at
the level of my neck or throat, and if I lean far
forward it gradually rises to my mouth

/ walked or half-walked, half-floated,
toward the cliffs. The rocks were almost bare in
this bay, like those between tides, and the
multitudes of lesser aquatic creatures were
concealed beneath them. The water was quiet,
and between surges was often perfectly clear, so
that I could see plainly the cliffs rising high in air
above that narrow straight line which marked the
division between the two kingdoms. I went as far
as my hose tether would permit and reached a
boulder on which, the day before, at low tide, I
had sat comfortably in the clear, cool air of the
upper world.

Turning back, I saw that I had become a
Pied Piper of sorts, leading a host of fish which
followed in my train. The sun was out now in full
strength and no fish, however strange and
unknown to me, could hold my eyes from the
marvel of distance. As I walked toward the cliffs I
had also worked a little toward the east and the
view I had, as I turned, was of another slope than
that over which I had come. The bottom thus far
was not wholly unlike the cliff above the water,
but before me now the slope fell away in a
manner which was beyond all experience — a
breath-stopping fall, down which one could not
topple headlong, but only roll and slide slowly,
to be overcome, not by swift speed of descent or
smashing blow, but by a far more terrible slow
increase of pressure of the invisible medium,
whose very surface film is death to us. ...

My range of vision was perhaps 50 feet in
every direction, but for all I could tell it might
have been 50 feet or 50 miles. The sun's rays
filtered down as though through the most
marvelous cathedral ever imagined — intangible,
oblique rays which the eye could perceive but no

lip describe. With distance, these became more
and more luminous, more wondrously brilliant,
until rocks died away in a veritable purple glory.
No sunset, no mist on distant mountains that I
have seen, could compare with this. One had to
sit quietly and absorb these beauties before one
could remember to be an ichthyologist.

As I was revelling in pure sensuous delight
at this color of colors, a small object appeared in
mid-water close to my little glass window, and
was instantly obscured by half a dozen little fish
which darted about it, some actually flicking my
helmet with their tails, lust as I saw that the
suspended object was a baited hook, a baby
scarlet snapper snatched at it, darted downward,
and was at once drawn up into the boat. As I
looked after it an idea came to me and I followed
the snapper upward by way of the ladder. When
the helmet was lifted off and I could speak, I
expressed my wants, and descended again. Soon
there fell slowly at my feet a small stone to which
was tied a juicy and scarcely dead crab. I picked
this up, waved it back and forth so as to scatter
the impelling incense of its body and as if by
magic, from behind me, from crevices upon
which I was seated, seemingly materializing from
the clear water, came fish and fish and fish. . . .

. . . Adam-like, I had to give them all
temporary names, until I could identify them, or
christen them with my own binomial terms. It
was long before I could disentangle individual
characteristics from the whirling mass. The first
four fishes rushed for the bait ... so (hat until I
could shut my mind to the abstract marvel of it
and my eyes to the kaleidoscopic, hypnotic
effect, ichthyology gained little of specific factual
contribution. Within three minutes from the time
when the crab first fell into my hand, I had 500
fish swirling around my crab and hand and head.
Similes failed. I thought of the hosts of yellow
butterflies I have seen fluttering at arm's length
on Boom-boom Point; I thought of the maze of
wings of the pigeons of St. Mark's, but no
memory of the upper world was in place here,—
this was a wholly new thing.

Acknowledgment

Reprinted by special arrangement with G. P. Putnam's Sons
from the book The Arcturus Adventure, Copyright 1926 by
William Beebe.

aegun quite late. The first attempts to harvest
•esources from these waters through diving were
issociated with the spiny lobsters — the red,
aanulirus penicillatus, and the blue, P. gracilis. It is
lommon knowledge that one used to be able to

walk along the rocky shore of the Galapagos and see
the antennae of these animals just below or even
above the surface of the sea, and that bags of them
were gathered by native residents with little effort.
During the 1960s, however, interest began in

Much has yet to be done to ensure a future for the islands'
spiney lobsters, such as the blue variety shown here.

a commercial catch. Divers captured the lobsters
using face masks alone. An original plan called for
the crustaceans to be stored in a large steel trap and
fed on fish until an airplane equipped with dry ice
came to collect them. There were reportedly about
10,000 lobsters in this trap at one time! The event
failed, though, due to poor arrangements, and the
lobsters were released, with 25 percent of them lost
as the result of crushing.

Nonetheless, the lobster business developed
rapidly with the arrival of freezer ships such as the
Villamil and Agrimar Chile. These large vessels
collected lobsters from a local, land-based freezer
plant, and also had their own divers. By now, swim
fins and snorkels had been added to the gear, and
some divers were starting to use wet suits. The
vessels operated as motherships to a flotilla of small
dinghies and greatly accelerated the exploitation of
this resource. On a round-trip from Guayaquil of 40
days duration, with 10 divers, 4,000 to 5,000 pounds
of lobster tails were extracted. The price in these
early days was about 9 U.S. cents per pound.

Both traps and bottom nets were eventually
experimented with, but both failed. The first traps
were constructed of wood, but these were
destroyed by sea lions who, we assume, wished to
get at the lobsters. Later, steel traps were
constructed. Although initially successful, they were
never used again, partly because of lack of
experimentation with baits. It was still cheaper to
dive for the lobster, and it was felt that people could
not be trusted to leave other people's traps alone!
Bottom nets, although quite effective in catching
lobsters, were also destroyed by sharks and sea lions,
so this experiment too was abandoned.

I suspect that SCUBA was first seen in the
Galapagos during the 1960s, when some work was
done with it in catching lobster. However, this failed
also, since the divers found that the bulk of the
equipment restricted their movement, making it
difficult to pursue their prey.

In the early 1970s, a vessel named the
Marisita arrived and began the use of surface-
supplied air instead of SCUBA. This method, which
is still utilized today, consists of the use of a normal
regulator supplied with air from a compressor carried
in a small dinghy equipped with an outboard motor.

Normally two divers operate down to a depth of
about 10 to 12 meters. Hand spears are still the
weapon of choice, as lobsters are a wary, spiney
prey. The spears also serve as protection from large
moray eels which at certain islands such as
Culpepper and Wenman, have been known to
follow divers over the reef. This equipment allows
the divers to penetrate deep into caves and recesses
along rocky coasts without the bulk of SCUBA. It is a
dangerous operation, however, and several divers
have been killed in recent years. The gear is often in
poor condition and no safety standards exist in the
relentless search for this profitable crustacean.

The end of the larger vessels came recently, in
the late 1970s and early 1980s, mainly because of
the unwillingness of the boat owners to pay their
men a steady wage, so as to guarantee them some
money during slack periods, or to give them social
security benefits. As it was, the divers were paid
solely on their catch. Toward the end of this period,
a diver received about 30 U.S. cents per pound of
lobster tail. This could be profitable, but, given the
poor visibility and dangerous coastal surf which
occur periodically, the divers often suffered physical
injuries from overextending themselves when
conditions were good.

Finally the divers, noting the rapidly rising
price of lobster, deserted their old masters and came
to the Galapagos with their own small vessels,
carrying surface compressors and refrigerators, and
now receiving up to 700 sucres, or about 5 U.S.
dollars, per pound of lobster tail. One cannot
criticize people for trying to make a living. However,
in the days of the large freezer ships, the divers
fished by day alone, whereas now they fish both day
and night, catching lobster in their daytime refuges
as well as when they are feeding at night. There is
still a certain amount of free diving (masks, fins, and
snorkel) done for lobster, too, but this is mostly by
local tourist boats to feed their passengers.

Black Coral

Although SCUBA may not have been popular for
lobster fishing, a new resource was discovered that
made SCUBA diving profitable: black coral,
Antipathes panamensis. Black coral was known to
exist here for many years. A local resident, Robert
Schiess, told me that, while he was fishing for
bacalao (a grouper) in the late 1940s, large clumps of
black coral were often drawn up — and thrown back
with a curse! If only he had known.

Today the extraction of black coral is an
expanding industry, bringing a price of about 2,000
to 2,500 sucres (about 16 U.S. dollars) per pound.
Remarkably beautiful when polished, the coral is
often made into jewelry or other ornaments. It is
difficult to enter a shop without seeing cases of it for
sale. Consequently, there is little hesitation by coral
gatherers or lobster fishermen to maximize the
harvesting of this material.

There is no doubt that both these resources,
lobster and black coral, have been exploited without
control. At one stage I was informed by the
Superintendent of the Galapagos National Park
Service that about 2,000 pounds of lobster tail a

week were being exported by air from the
Galapagos. Since 1971, it has been illegal to take
lobsters over 25 centimeters in length or having a tail
weight of more than 180 grams, as well as gravid
females. But the use of hand spears often mortally
wounds the lobster before the diver gets close
enough to see whether the creature is undersized or
a gravid female. In 1985, a moratorium on lobster
catching of any sort for four months a year
(December-January and June-July) was imposed for
further protection. But to judge whether these laws
are obeyed is very difficult. There is so little control
that there are almost no details available. During an
inspection of the lobster vessel, El Salvador, 20
percent of the lobsters were discovered to be
undersized. The only inspectorate is in San Cristobel,
and as most of the lobster is exported from the
airport of Santa Cruz, enforcement is virtually
nonexistent.

The lobster population must vary naturally
from year to year depending on oceanographic
conditions, but as far as I am aware, no study has
been carried out to understand the dynamics of the
population. It is unclear as to whether the juvenile
population originates solely from Galapagos adults,
or from an influx of larval forms from the central
Pacific area as well. But that the population has
suffered there can be no doubt. From all sides,
reports come in as to the shortage of lobsters
available.

It is difficult to believe that the lobster
population in the Galapagos region will become
extinct, since there are many areas that are
extremely difficult to fish, but it does seem a shame
that a locally useful industry can be spoiled by short-
term greed. Clearly, more must be done to ensure a
future for the lobster and the lobster industry.

The black coral industry goes on almost
completely unchecked. As with the lobster, there are
few statistics available, but damage is surely being
done. The coral's growth rate is believed to be slow.
The only brake applied to the industry is through the
National Park Service where guides try to explain the
problems and ask people not to bag the product.
Judging from the vast amount of sale, this is not
enough.

Obviously, with the introduction of the
Marine Resources Reserve, we expect the
implementation of laws to control the exploitation of
many of the local species, including lobsters and
black coral. One would expect large areas to be set
aside as scientific reserves with permission to dive
only (no collecting at all) with carefully controlled
anchor sites and/or mooring buoys. (At many
anchorage sites, diving under the vessels is like
diving in a garbage dump, with old filters, plastic
buckets, and so on.) Some areas will have to be set
aside for traditional lobstering and fishing, and
perhaps licensed coral dives, but preferably the coral
business could be phased out altogether.

However, the enforcement of such laws will
always be extremely difficult because of a lack of
money for transport, radios, and other inspecting
equipment, and a lack of wages for park personnel.
The best park employees often leave the service for

Black coral, Antipathes galapagensis. (Photo by T. M. Rioux,
WHOI)

the more lucrative wages offered by the tour trade as
guides or captains. Effective control will be
expensive but is essential to the future of the
region's marine resources.

Sport Diving

A second major use of diving equipment in the
islands is also profit-motivated: the tourist trade,
which utilizes both snorkel and SCUBA equipment.
The very latest in designs can be seen here (although
there are no dive shops on the islands). This modern
gear, on the one hand, is very safe, but its
introduction has accelerated the destruction of coral
beds. Sport diving started about 1969 or 1970.
Although it began slowly, it has now become a major
industry, with a number of tour operators offering
diving as an optional extra to their normal terrestrial
activities. Diving around these islands is certainly
fascinating and unique — one can be lucky enough to
dive with penguins and flightless cormorants, with
marine iguana, with sea lions and fur seals, with
sharks and large grouper and snapper, rays and large
eels.

There are walls of fish to be seen, yellow-
tailed surgeon fish, goat fish, grunts, Creole fish and
white-banded angel fish. Ecologically the fish are
very interesting, some with their origin in the warm
tropical waters to the north, others from the cold
waters to the south, and more still from the central
Pacific area. There are corals of several species,
including black coral colonies, whose beautiful
waving fronds look like feathery submarine bushes.
There are many habitats: steep walls, rocky bottoms,
sand bottoms, caves, and areas rich in algae where
marine iguanas and beds of garden eels may be seen
feeding.

Restrictions Necessary

For commercial and sport diving to coexist, control is
vital. Commercial divers need laws restricting them
to protect their own future and to prevent them for
despoiling areas where sport diving is active. Sport
divers need laws to stop them from doing ecological
damage by removing such objects as shells or
endangering other divers and upsetting ecological
balances by spear fishing. The philosophy of

protecting areas for future generations must apply to
our submarine world as much as to our terrestrial
one.

The Galapagos are not an easy place to dive.
The area is well known for its large animals, such as
sea lions, sharks, and eels; the waters are often cold
(16 to 18 degrees Celsius); sharp thermoclines exist;
visibility can be excellent or very poor; and in some
regions currents are very strong (2 to 4 knots). It is
not an area for the inexperienced amateur diver.
One needs confidence in one's equipment and
confidence in one's self.

Remarkably enough, there has been no major
accident among sport divers yet, be it from animal
attack or failure or misuse of equipment. The sharks
that are seen are relatively quiet, although several
people have had close encounters with reef
whitetips, and some have even been bumped by
them. Once I dived through a cave and, on coming
out the other side, found myself in the midst of a
group of about 50 hammerhead sharks.
Transformation into a rock is difficult at that point.

There is no doubt that procedures should be
tightened. There is virtually no emergency help in
the islands. Metropolitan Touring (the area's largest
tour company) provides its yachts with an oxygen
supply, but the quality of the oxygen is unknown.
The nearest immediate help that can be counted on
is Panama, from where the Howard Air Force Base
will send an airplane free of charge. However, use of
the decompression chamber in the Canal Zone
carries a minimum charge of about 3,000 U.S.
dollars.

Recently, David Balfour, the manager of
Metropolitan Touring's office in the Galapagos, gave
me some points which he considers indispensable
for diving groups:

/) Croups should be exclusively oriented to
diving;

2) Each diver should provide a Certificate of
Health and a Certificate of Diving
Competency (although many of these seem to
be of doubtful use);

3) The Galapagos tour operator should provide a
guide who is a qualified diver to be
responsible for and accompany each group of
divers;

4) Only tanks, weights, and a compressor are
provided in the islands. Regulators and all
other equipment are the total responsibility of
the diver.

I would add that buddy diving must be the
rule. When 10 divers are in the water, it is
impossible for one person such as the guide to
account for all people at all times. In addition, no
decompression diving should be attempted.

These controls may be possible to impose on
tour groups, but for the commercial diver, it is
difficult to see in the near future any realistic
solution. Their business is profit-motivated and they
will take risks.

Scientific Diving

The most recent development in diving in the

Galapagos was the arrival in 1986 of a submersible
on board the vessel Seward Johnson, owned by the
Harbor Branch Foundation. The work of the
scientists on board involved collecting many forms of
invertebrates and algae in the hope of discovering
chemicals which may help in the control of various
human ailments, such as tumors and in the
production of antibiotics (see article page 69).

Securing a Future

The archipelago thus may be economically
important to its residents, aesthetically pleasing to its
divers, and provide medicines yet unknown. Diving
in the Galapagos has grown rapidly and is basically
profit-motivated. To secure a future, it is now
necessary to clarify the situation and to impose and
encourage conditions that will help to conserve the
environment, yet at the same time allow some to
make a living and others to enjoy a still magnificent
undersea environment.

Godfrey Merlen, an Englishman, is a Naturalist Guide, marine
artist, and underwater photographer working in the
Galapagos. He recently completed a guidebook to
Galapagos fishes, which is currently in press.

Suggested Readings

Beebe, C. W. 1926. The Arcturus Adventure. New York: C. P.

Putnam's Sons.
Clynn, P. W., and C. M. Wellington. 1983. Corals and Coral Reefs of

the Galapagos Islands. Berkeley: University of California Press.
Martinez, P., and C. Robinson. 1984. Investigaciones sobre las

explotacion del coral negro (Antipathes panamensis) en las Islas

Galapagos, Ecuador. Boletin Cientifico y Jecnico del Institute

Nacional de Pesca, Vol. 6, No. 3, pp. 107-123. Guayaquil,

Ecuador.
Reck, G. K. 1984. La pesca de langosta en las Islas Galapagos, 1974-

1979. Boletin Cientifico y Jecnico del Institute Nacional de Pesca,

Vol. 6, No. 3, pp. 49-77. Guayaquil, Ecuador.

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Galapagos white tip shark and turtle. (Photo courtesy of Feodor Pitcairn, © 1984, from Hidden Seascapes, published by Little,
Brown & Co.)

A squadron of Galapagos manta rays. (Photo courtesy of Feodor Pitcairn,
Brown & Co.)

1984 from Hidden Seascapes, published by Little,

The Fishes of The
Galapagos Islands

by John E. McCosker

IVlore than a century and a half ago, a young
naturalist named Charles Darwin visited the
Galapagos Islands. Like many of his modern
counterparts in marine biology he suffered from
mat de mer, or seasickness, and was overjoyed at
the sight of land. Being a curious British naturalist,
he dropped a line overboard to sample the strange
fishes. The 15 specimens he returned to England
with were all subsequently described as new
species, largely a reflection of the existing state of
the art in ichthyology at the time.

Were the aquatic flora and fauna of the
mainland and islands to the west better known at
that time, he could have based his theories of The
Origin of Species by Means of Natural Selection . . .
on the fish, or ichthyofauna, alone. The endemism
(the fact that they live nowhere else) of the
Galapagos nearshore fish parallels that of the
terrestrial tortoises and finches — whose ancestors
found themselves in offshore isolation, then
mutated, adapted, were selected upon, and
evolved to become textbook examples of the
process of evolution.

Oceanographic Factors

The unique nature of the fish in Galapagos waters
stems from their complex geographic and
oceanographic environments. These have been
well summarized by Guy T. Houvenaghel, an
oceanographer at the Universite Libre de Bruxelles,
and elsewhere in this issue.

The archipelago rises abruptly from the
intersection of the Cocos and the Carnegie
submarine ridges (see map page 13), forming a
platform separated from the mainland by deep
oceanic water no shallower than 1,300 meters. The
closest inhabitable shallow reefs are those of the
island of Malpelo, a small rock 435 kilometers to
the northeast. These reefs probably serve as
stepping-stones for certain nearshore species to or
from the mainland.

In the past, geologists and paleontologists
heatedly argued the age of the Galapagos. The

Above, the Galapagos batfish, Ogcocephalus darwini, a
species endemic to the archipelago, resting over a sand
bottom at 10 meters. (Photo by C. Roessler)

former favored youth, perhaps no older than 3+
million years ago (MYA), and the latter, an age as
far back as the Miocene (10 to 14 MYA). Recent
paleontological evidence indicates that adaptive
radiation of Galapagos terrestrial species occurred
within the past 3 to 4 million years, apparently
vindicating the geologists. Shallow water reefs may
have existed several million years prior to the
terrestrial emergence, however.

The archipelago consists of several
emerging, steep-sided mountaintops of huge
submarine volcanoes. The water barriers and/or the
distance between the mountaintops has allowed
the speciation of animals on shore, but this is not
enough to stop the distribution of marine larvae.
The endemic fishes are typically found at each of
the islands possessing appropriate habitats.

A typical shore profile slopes off along sand
and shell bottoms, with rich algal development in
cooler waters and sparse coral development along
the warmer shores. True coral reefs are absent, in
that hermatypic (reef-building) corals cannot
tolerate the cool temperatures of the islands. The
northernmost islands (Darwin and Wolf) and banks
possess the greatest abundance of coral and Indo-
Pacific fauna.

Several currents meet and mix at the
Galapagos. The islands are washed by the cold,
salty waters of the Humboldt or Peru Current from
the southeast and the warmer, fresher Panamic
surface water from the east. From the west comes
the Equatorial Undercurrent which, during non-El
Nino years, is a stable water mass encountering the
northwest edge of the archipelago and bringing
with it larval and adult marine organisms from the
central Pacific. During the extreme El Nino event of

1982/83, much of the shallow nearshore
environment was markedly affected, including the
decimation of many cold-dependent species of
fishes, invertebrates, algae, and the marine iguanas
(see article page 54). This event also brought
numerous central Pacific tropical species to the
islands.

It is likely that the extreme temperatures and
ephemeral conditions associated with extreme
El Ninos result in the transient appearance of
certain tropical Pacific species. This explains the
fact that several Galapagos records are based on
but a single specimen. The sharpnose pufferfish,
Canthigaster amboinensis, and the sailfin leaffish,
Taenianotus triacanthus, are examples of this
phenomenon. Other species have been sighted but
not collected, such as Klein's butterflyfish,
Chaetodon kleinii. Such species were perhaps
unable to colonize for lack of hardy larvae, or have
become extinct once water temperatures became
lower.

The strong equatorial surface winds create
upwelling conditions along the western shores of
several islands (primarily Fernandina and Isabela).
That difference of 5 to 10 degrees Celsius results in
lush algal growth, and the presence of fishes and
invertebrates typical of the more temperate
Peruvian-Chilean flora and fauna. Observations of
parrotfish behavior by ichthyologists Richard H.
Rosenblatt of the Scripps Institution of
Oceanography and Edmund S. Hobson of the
U.S. Fish and Wildlife Service attest to this
phenomenon. During February 1967, an inter-El
Nino year, they recorded a near surface
temperature of 26.7 degrees Celsius along the
northern shore of Hood Island. The southern shore

The Galapagos four-eyed blenny, Dialommus fuscus, on a terrestrial sojourn in search of food. (Photo by D. /. H. Phillips)

temperature was 21.7 degrees Celsius at the same
depth. Only a single parrotfish species, the
loosetooth parrotfish, Nicholsina denticulata, was
seen along the southern shore. But four species of
tropical parrotfishes were found along the other
side of this rather small island. Thus, the upwelling
phenomenon allows the presence of two rather
distinct faunas on either side of an island, and
significantly increases the faunal complexity of the
Galapagos.

Ichthyological History

Early explorers to the Galapagos collected a
few specimens of fishes, but it was Darwin that
made the largest early sampling. His 15 specimens
were described by his friend and fellow-naturalist,
the Reverend Leonard jenyns, in 1842. Included
among those specimens was the Galapagos
sheephead wrasse, which Jenyns named Cossyphus
darwini in honor of its collector (see photo page 29
and drawing page 77).

Thirteen years later, French ichthyologist
Achille Valenciennes reported on the fishes
collected by an expedition aboard the frigate
La Venus in 1838. He described 13 new species,
and was the first to recognize the unique
distribution of the insular fishes. Although largely
ignorant of the eastern Pacific ichthyofauna, he was
able to observe that the Galapagos fishes were
more closely related to those of "Atlantic America"
than to those of the Indian Ocean.

The first American expedition to the

Guinea fowl pufferfish, Arothron meleagris, in its normal
coloration, a species common to the tropical Indo-Pacific.
(Photo by R. Ames)

Galapagos was led in 1873 by Professor Louis
Agassiz of Harvard University, accompanied by
zoologists Franz Steindachner of Vienna and Count
Louis Francois de Portales aboard the Hassler. This
was followed by expeditions aboard the U.S.

Odd Fish of the Islands

he unique oceanographic conditions
of the Galapagos have acted on the shorefish
fauna to allow the evolution of several
remarkable species. For example, only at the
Galapagos can one see the marine iguana,
Amblyrhynchus cristatus (see article page 54),
making its daily procession from its volcanic
perch to the sea, in search of the red algae on
which it dines. In the opposite direction, one
may observe the Galapagos tour-eyed blenny,
Dialommus fuscus (see photo page 30), a
diminutive, mottled fish, whip-sawing its way up
the shore in search of its diet of insects and
shore crabs.

The terrestrial sojourns of the blenny may
take it as far as 30 meters from the sea, far from
its normal predators, the groupers and snappers.
Dialommus has had to adapt to the problems of
vision, locomotion, and respiration out of water.
Its eye surfaces are most remarkable, in that the
corneas are laterally flattened and meet at an
angle of approximately 100 degrees along the
vertical midline. Studies by Richard H.
Rosenblatt and leffery Graham of the Scripps
Institution of Oceanography show that such fish
avoid myopia by adjusting the refraction of light
incident on the cornea. In this way, the lens will
produce a clearly focussed image on the retina.
Gill modification, such as thickening and
filament enlargement, and behavioral
adaptations allow the fish to breathe air for as
long as two hours.

As a diver or snorkler, one is visually
treated in the Galapagos to the harlequin
wrasse, Bodianus eclancheri, a lovely
sheephead relative that exists in a myriad of
color combinations. Its variable splotches of

orange, crimson, black and white are
reminiscent of the patterning of a Koi carp. This
apparently uncontrolled harlequin coloration
was studied by Steven Hoffman, then at the
University of California at Santa Barbara. He
explained it in terms of selective pressures in the
Galapagos.

The harlequin wrasse is a sequentially
protogynous hermaphrodite (changing sex from
female to male), but differs from related species
in not being sexually dimorphic. It has
abnormally large gonads, and the sexes are
equally active. This seemingly unusual behavior
is explained by Hoffman and his professor,
Robert Warner. They theorize it to be a
response to the predation pressures of
Galapagos sea lions, Zalophus californianus
woilebaeki, and Galapagos reef sharks,
Carcharhinus galapagensis. To avoid the
extraordinary predation at the Galapagos, males
of this species have increased their feeding and
decreased their reproductive activities. These
behaviors are quite unlike those of their
mainland relatives.

As a final example, the Galapagos batfish,
Ogcocephalus darwini (see photo page 28), is
notable in that it demonstrates the influence
that the absence of predators plays in isolated
situations. Batfishes are small, sedentary, sand
and mud bottom fishes. Along the eastern
Pacific mainland, batfishes are rarely seen in
shallow water and are probably limited in their
range by the activity of many predators. The
Galapagos batfish however, is common below
10 meters, and is so incautious as to be hand-
captured by the SGUBA diver.

—JEM

Fisheries vessel Albatross, and the extensive
collections of Stanford University ichthyologists
Edmund Heller and Robert Snodgrass aboard the
sealing schooner Julia E. Whalen. Their activity, as
well as the specimens collected by New Bedford
whalers coming in for provisions at the Galapagos,
brought the fish tally to 128 species by the end of
the century.

In the early years of the 20th century,
several wealthy Americans brought museum-
associated scientists to the Galapagos aboard their
fashionable yachts. William Beebe, the celebrated
ichthyologist of the New York Zoological Society,
visited the Galapagos aboard the Noma and the
Arcturus. His accounts were published in both
popular volumes (see page 22) and scientific
papers. Other visits included Captain Allan
Hancock's Oaxaca and the Ve/eros /-///. Vanderbilt's
Cress/da also made the voyage.

More extensive collecting efforts, aided by

modern SCUBA techniques and rotenone
ichthyocides (plant-derived fish poisons), were the
1964 Galapagos International Scientific Project, the
1977 California Academy of Sciences, and the 1984
Los Angeles County Museum expeditions. The
1977 expedition that I led discovered several new
species and new records by diving to more than 60
meters. The 1984 expedition, led by ichthyologists
Daniel Cohen, Robert Lavenberg, and Jack Grove
was fortuitous in that many important observations
and discoveries were made in the wake of the
extreme El Nino event.

The curious nature of the Galapagos and its
collectors has resulted in some unique captures.
For example, a National Geographic team
photographed a strange sailfin leaffish, Taenianotus
triacanthus, in shallow water at Albany Island in
1978. It has never been seen since. Andre De Roy,
a longtime resident of the Galapagos and avid shell
collector, presented this author with the first and

only known specimens of a highfinned
ateleopodid, Guentherus a/t/Ve/a, and a smooth
stargazer, Kathetostoma averruncus. He collected
the stargazer by dragging a coffee can dredge in
600 meters behind his boat. In 1980, I dropped a
line to the shallow 7-meter bottom while at anchor
aboard the Encantada off Punta Espinosa. The result
was the capture of a large female spotted
houndshark, Triakis maculata, the only known
specimen. These unique captures reflect the
apparent scarcity of certain species, the lack of
collections, and the ephemeral nature of some
species occasionally arriving at the Galapagos.

Faunal Complexity

The number of fish species in the Galapagos is
large when compared to that of many other
tropical eastern Pacific islands. The size of this
number is due to the diversity of habitats, the large
area of the archipelago, and the various currents
sweeping the islands. The currents bring larvae of
three quite different biogeographical regions. My
colleague Rosenblatt and I now consider it to
contain 307 species, representing 92 families
(Easter Island to the south has only 110 species, but
the Hawaiian Islands have more than 470
according to ichthyologist John E. Randall of
Honolulu's Bishop Museum).

The high faunal complexity of the Galapagos
is directly related to its variety of habitats. Some
examples are rocky shore, sand bottom, and
mangrove environments. But the near absence of
muddy bottom communities puts an upper limit on
this complexity. Another factor limiting the number
of Galapagos fish species is the difficulty that many
mainland shorefish groups face in crossing 1,000
kilometers of ocean.

Ichthyologists consider the Galapagos fish
fauna to be a distinct subunit of the "Panamic
province"; with other elements from the Chilean (4
percent), the western Pacific (14 percent), and the
eastern Atlantic provinces in addition to endemic
species. Nearly 60 percent of the fishes are shared
with the eastern tropical Pacific mainland. About 8
percent are worldwide in the tropics, or
pantropical, such as the oceanic sharks (whitetips,
whale sharks, and hammerheads), manta rays,
pufferfishes, tuna, and dolphinfishes. Certain large
wrasses and groupers are common to Peru and
Chile, and four species, excluding the pantropicals,
are common to the western Atlantic.

As previously mentioned, the high degree of
endemism of Galapagos shorefishes is particularly
instructive. At least 51 species, or 17 percent, are
unique to the Galapagos, a condition similar to that
of the Hawaiian Islands and other oceanic
outposts. Among the endemics, a group of 7
species is found only at the Galapagos, Malpelo,
and Cocos Island to the north.

Marine fish endemism is best explained by
the vagility, or transportability of the larva and/or
adult, and the duration of larval life of each
species. Those fishes that are strong open ocean,
or pelagic, swimmers, such as the jacks or tunas,
have no difficulty in crossing the 1,000 kilometers

between the mainland and the islands. Other
species with protracted larval stages well-suited to
pelagic life are certain groupers, blennies,
damselfishes, and the almost-invisible moray eel
larvae. Adult or larval forms that inhabit floating
detritus, such as wrasses and certain blennies, are
also able to make the crossing. This continual
opportunity for genetic interchange reduces the
opportunity for species to evolve.

Endemism, however, favors those forms with
short larval lives unsuited to pelagic transport.
Examples are croakers and grunts, which can then
speciate without continual genetic swamping (but
only if they fortuitously arrive in the first place).
The endemic Galapagos species are usually well
differentiated from their closest relatives.

The majority of endemics are most closely
related to Panamic species, as one might expect
from their proximity to the New World, and the
great distance separating them from the western
Pacific. Typical are the chaenopsid pike blennies,
clinid klipfishes, and stargazers — all related to New
World species. Four species are related to forms
from Peru and Chile. Others do not fit the pattern,
such as a burrowing snake eel, Callechelys
galapagensis, related to a Hawaiian form; and a
porgy, Archosargus portalesii, related only to
western Atlantic forms.

Future Research

Although the nearshore fish fauna of the
Galapagos, within the limits of comprehensive
SCUBA collecting, is now fairly well-known,
surprises remain. It is difficult to sample the craggy
bottom below 50 meters, thus hindering adequate
bottom net-trawling. This means that new species
and new records remain to be discovered.
Improvements in diving techniques will allow
ichthyologists to collect in deep reef habitats.

Several long-standing handicaps, hindering
Galapagos ichthyologists from becoming involved
in issues of concern to their terrestrial counterparts,
are about to be overcome. We will soon be able to
pinpoint the geologic age of the submarine
environment, and the survey of fishes is becoming
ever more complete. This information will be used
in comparing the evolutionary rates of marine
animals to those of the finches and tortoises. We
also need to explain and predict the filling of
niches, or ecological saturation. In the Galapagos
this information is crucial to fisheries biologists
concerned with the introduction of exotic species.
Finally, we are just beginning to understand the
significance of the infrequent genetic influx
allowed by rare climatic events, such as the
extreme El Ninos. Results of these projects will
undoubtedly answer some questions, but pose
others for future Galapagos ichthyophiles.

lohn E. McCosker is Director of the Steinhart Aquarium of
the California Academy of Sciences in San Francisco. His
research interests include the fishes of the Galapagos, and
the behavior of the white shark, penguins, and coelacanths.

Marine Biological Research

in the Galapagos:

Past, Present, and Future

by Henk W. Kasteleijn

V V hile the land-based biology of the Galapagos is
familiar to many people, the islands' marine
biology is much more poorly understood. This is
unfortunate because many animals, such as the
marine iguanas and seabirds, depend heavily on
the marine environment. In the last 10 years, the
pace of research into the Galapagos' marine
biology has been increasing, but the first
observations on marine life were made during the
discovery of the islands themselves.

The Early Years

When Fray Tomas de Berlanga, Bishop of Panama,
discovered the Galapagos in 1535, he wrote a long
letter to his king in Spain. He described not only
the islands and their terrestrial life, but also some of
the marine creatures. He mentioned the seals and
one true marine animal, the green turtle, Che/on/a
mydas.

Ambrose Cowley, a buccaneer on board the
Bachelor's Delight with Captain Cook, remarked in
his diary of June 1684 on the abundance of fish life
in the Galapagos. He also wrote that green turtles
(see box page 34) were so plentiful "that though
wee were about 200 soules yett wee killed every
morning on the Bay as many as served us all day
the whole time of our abode there and might have
kill'd many more. Though they differr in nature
from the West India Turtle yett are very sweet
wholesome meate." In addition to the green turtle,
he mentioned the presence of the hawksbill and
the loggerhead, which he described as "neither
good flesh nor shell."

Another buccaneer, Ravennau de Lussan,
made a very strange observation in 1684: "These
adjacent waters are also full of fishes, that come up
to die on land." Possibly a school of fish was
cornered and driven ashore by sharks. Although
the cause of such incidents remains unclear, one
was recently filmed in the Galapagos by Dieter
Plage, of Survival Anglia Films.

The large number of whales originally
frequenting the Galapagos seas was first described
in 1 700, by Ensign Le Sieur Villefort, a French

seaman aboard the frigate Philippeaux. The first
organized reconnaissance of whales in the vicinity,
however, was conducted in 1793-94 by Captain
James Colnett of the Royal Navy. On this voyage,
for the benefit of the British whaling industry, he
also made the first workable chart of the
Galapagos.

Aboard the Rattler, Colnett was mainly
interested in the exploitable spermaceti whale,
Physeter catodon, which he mostly saw in groups of
cows with their calves. Other large whales that he
found were the humpback whale and the fin
whale; but he also mentioned large schools of
porpoises, killer whales, arid "blackfish" (small,
dark, blunt-headed whales such as the melon-
headed whale, pygmy killer whale, false killer
whale, and short-finned pilot whale).

Captain Colnett also caught "great numbers
of large Cod," within a short time at Kicker Rock,
off San Cristobal Island. These were probably the
very abundant, endemic bacalao or yellow grouper,
Mycteroperca olfax. Other species caught on the
1794 voyage were albacore, mullet, devilfish or
manta ray, and bonita.

"Sharks were in great abundance," Colnett
noted. He also saw squid, rocks covered with
crabs, and a "few small wilks and winkles." Finally,
he wrote in his journal that "A large quantity of
dead shells, of various kinds, were washed upon
the beach, all of which were familiar to me; among
the rest were the shells of large cray-fish, but we
never caught any of them alive."

One of the most famous whalers to visit the
Galapagos was the American novelist Herman
Melville, who was an able seaman aboard the
whaling ship Acushnet. He chronicled his
impressions of the archipelago in his short novel,
The Encantadas (see article page 93).

For the Galapagos, the 19th century can be
characterized as the century of over-exploitation.
The whaling industry was at its peak from about
1790 until 1840. During this time, the whalers took
blackfish, and many terrestrial and marine animals.
But when the sperm whale stock in Galapagos
waters was depleted, the collapse of-the whaling
industry in this part of the world was at hand.

Galapagos Sea Turtles

I V/ar/ne turtles are perhaps the most mysterious
form of all Galapagos wildlife. They remain below
the waves for nearly their entire lives, hidden
from the marine biologist, who's best chance to
study them is during the relatively brief time they
spend on land, in nesting and hatching.

The east Pacific green turtle, Chelonia
mydas agassizii, ;s by far the best-represented
species of sea turtle in Galapagos waters. But
Derek Green of the Charles Darwin Research
Station admits that it is impossible to give a good
estimate of their population. Green conducted an
extensive study of their nesting behavior from
1970 to 1979, and was able to tag nearly 4,000
females and 10 males during the study.

This particular species of marine turtle,
averaging about 50 to 100 kilograms in weight
(some large individuals can approach the weight
of the Galapagos giant tortoises), is known locally
as la tortuga negra, or "black turtle." The
archipelago's inhabitants also speak of a tortuga
amarilla, or "yellow turtle," but it is very rarely
seen. Biologists believe it to be a sterile mutant of
Chelonia mydas agassizii.

Two other species of sea turtles are
sometimes sighted nearby, but neither nest on
the islands. Green says the Indo-Pacific hawksbill,
Eretmochelys imbricata, is encountered
occasionally, and that the Pacific leatherback,
Dermochelys coriacea, was seen in the area only
three times prior to 1 979.

The Galapagos are probably the most
important nesting area in the east-central Pacific
for the tortuga negra, according to Green (see
map page 36). The most popular beach in the
archipelago for this activity is Quinta Playa, on
the southeast shore of Isabela. As many as 45
females laid their eggs there one night in 1978.
That year, 610 nesting females were recorded on
that beach.

Despite the many eggs laid at Quinta
Playa, eggs laid at other beaches stand a much
better chance of hatching. Feral pigs and the
scarabeid beetle, Trox suberosus, prey on the
eggs, and both predators are prevalent on
Isabela. The beetles attack turtle nests on some of
the nearby beaches, but most of the other
important nesting beaches on the islands are safe
from them.

The pigs pose a greater threat throughout
the Galapagos. They seem to be able to identify
clutches of turtle eggs the way their relatives in
France locate truffles. At Espumilla beach, on
James Island, feral pigs were responsible for the
destruction of more than 98 percent of the eggs
laid there in 1979. The (lack of a) maternal
instinct in tortuga negra females does little to
help the situation, either. Darwin Station field
assistant Mario Hurtado reported a case where a
pig gobbled up the eggs even as the oblivious
female was producing them!

It is sometimes said that when females lay
their eggs, they return to the beach where they
themselves were hatched. If that is the case, in
the future there should be increased nesting on
beaches with less predation of eggs and hatch-
lings. Such beaches are Las Bachas on Santa
Cruz, and Las Salinas on Baltra Island.

One of the biggest mysteries concerning
the tortuga negra is their migration patterns.
Biologist Green believes that some of them are
present in the Galapagos year-round, based on
recaptures of tagged turtles. But he also reports
that individuals tagged in the archipelago have
been recovered from as far away as Costa Rica,
and Peru. He says that without more recapture
data it is impossible to know what proportion is
truly resident and what migratory.

The creation of the Galapagos Marine
Resources Reserve will probably have little effect
on sea turtle status within its boundaries. The
turtles have already been protected on the
beaches throughout the islands, where they are
most vulnerable; and since their flesh "is not
highly esteemed" by the local inhabitants, very
few have been taken from the sea in recent years.

Of course, the marine reserve can only
protect those turtles that stay within it, and all
three species found in the Galapagos are
exploited to some extent along the South
American coast. The relatively large population of
marine turtles protected by the reserve is of
international importance. As long as funds remain
available for biologists to continue their research,
the mysteries of the marine turtles will be steadily
unravelled.

— TMH

Continuing beyond the collapse of the
whaling industry was the fur seal hunt. But by the
end of the century the Galapagos fur seal,
Arctocephalus galapagoensis, was likewise chased
to the brink of extinction.

Of course, all the whalers and hunters

required food. C. H. Townsend, in an extensive
study of whalers' logbooks published in Zoo/og/ca,
estimated that more than 100,000 giant tortoises
were killed within a period of 30 years. Several of
the original 14 subspecies of tortoises were brought
close to extinction by this exploitation (see article

Marchena £

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Nesting Beaches

1 Quinta Playa

2 Bahia Barahona

3 Las Bachas

4 Las Salinas

5 Bartolome

6 Espumilla

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Principal nesting beaches of the east Pacific green
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page 86). The number of sea turtles that were
slaughtered may have been as large or even larger,
since they were good meat and more easily
accessible. There is no information, however, on
the effects of this exploitation on marine turtle
populations.

Early Scientific Expeditions

The first explorer making ichthyological collections
for study rather than consumption was Charles
Darwin. Although a connection between the
marine iguana and the sea had been noted before
he arrived, Darwin was the first to verify the link.
By examining their stomach contents, he learned
that marine iguanas eat algae. But he also was
interested in shells. He knew of the work of Hugh
Cuming, who collected in the Galapagos around
1829 aboard the Discoverer, and noted that more
than half of the 90 species he gathered were not
found anywhere else (see Darwin's Beagle account
page 72).

Robert FitzRoy, Captain of the Beagle, made
excellent charts of the Galapagos, which were in
use until 1942. At that time, the U. S. S. Bowditch
surveyed the islands and found that FitzRoy's
charts needed only slight improvement (see box
page 37).

Later Scientific Expeditions

Some of the important scientific expeditions to the
Galapagos of the late 19th and early 20th centuries
have been recounted elsewhere in this issue, and
included among them were the research voyages
of the U. S. Fish Commission's Albatross in 1888
and 1891. Louis Agassiz's son, Alexander,
accompanied the 1891 trip as chief naturalist.
Using new techniques, he did much to advance
the biological and geological knowledge of the
islands.

Alexander Agassiz employed systematic
dredging to assay the communities of jellyfish,
anemones, sea urchins, and starfish near the
islands. From his deep-sea work, he concluded that
the Galapagos must have had a volcanic origin,

The east Pacific green turtle, Che/on/a mydas agassizii,
the only marine turtle to nest in the Galapagos. (Photo
by D. J. H. Phillips)

since he found no topographical bottom continuity
between the archipelago and the Americas.

Rollo H. Beck had already made three
collecting trips to the Galapagos when he was
chosen to lead the California Academy of Sciences
Expedition of 1905-06. This was the longest and
most elaborate collecting expedition to reach the
Galapagos yet. They travelled aboard the schooner
/Academy, and spent a year and a day in the
archipelago. During their stay, they collected
recent as well as fossil (especially marine mollusk)
material.

Many other scientific expeditions travelled
to the islands, steadily increasing the knowledge of
Galapagos fishes and marine invertebrates. Some of
those of greatest interest with regard to marine
fauna are the St. George expedition of 1924 and
the Norwegian Zoological Expedition aboard the
Monsunen of 1925. Karl P. Schmidt of Chicago's
Field Museum of Natural History visited the islands
on the Illyria in the early part of 1929, as head of
scientific staff of the Cornelius Crane Pacific
Expedition. He returned to the Galapagos later that
same year on the Mary Pinchot as a member of the
Pinchot South Sea Expedition. Schmidt made some
interesting observations on marine iguanas on his
first voyage, and both expeditions resulted in the
description of new fish species.

Perhaps the most prominent among these
early scientists was William Beebe. His work is
described on page 22.

Millionaires

The wealthy readers of Beebe and Melville were
often inspired to view the wonders of these
enchanted islands for themselves, leading them to
mount expeditions aboard their private luxury
yachts. They usually carried naturalists who could
explain the unique fauna and flora they
encountered, and these naturalists were often able
to conduct research along the way. Some of these
millionaires were particularly interested in marine

Captain FitzRoy of H.M.S. Beagle

I he importance of Robert FitzRoy's visit to the
Galapagos 150 years ago has been completely
overshadowed by the fact that he was
accompanied by Charles Darwin. Darwin was
definitely the junior partner at that time, but
today FitzRoy is recalled almost solely on
account of his denunciation of the Origin of
Species a quarter of a century later.

This cause of dissension did not exist in
1835. FitzRoy was not then a serious student of
the Bible and a confirmed fundamentalist, nor
had Darwin become convinced of the
mutability of species. Despite profound
differences of temperament and political views,
these two brilliant young men (FitzRoy was
given command of H. M. S. Beagle at the age of
23) were still good friends when they got back
to England after sharing a cramped cabin in a
tiny ship for five years.

FitzRoy had a life-long interest in science
(it was entirely on his own initiative that he
invited a naturalist to be his guest on the
voyage) and he showed outstanding talent in his
own specialist fields. The object of the Beagle's
voyage was not to revolutionize biological
theory nor to provoke the greatest intellectual
debate of the century, but to chart the coasts,
chiefly in South America. FitzRoy did this with
remarkable skill. The buccaneer, Ambrose
Cowley, had made some rough sketches in
1684 and Captain James Colnett of H. M. S.
Rattler had improved on them in 1 793, but
FitzRoy's Galapagos charts are in a different
class and are barely distinguishable from those
in use today. The captain of the French ship, La
Genie, who came to survey the islands in 1846,
wrote of FitzRoy's achievement:

Nothing escaped the perspicacity of this
conscientious observer: the smallest
details are all indicated with really
astonishing precision and following his
drawing one can visualize in the most
accurate manner the shape of the coast.
Coming after him there is not even an
opportunity to glean.

/. R. Slevin, the historian of the California

Academy of Sciences' great research expedition,
wrote in 1959:

It is truly amazing that the modern chart of
the Galapagos made in 1942 by the U.S.S.
Bowditch equipped with every modern
device should so closely approximate the
survey made by Captain FitzRoy over a
hundred years before. His little vessel was
at the mercy of strong and uncertain
currents, together with deadly calms so
prevalent in those regions.

When he retired from active service in
1850, Admiral FitzRoy was elected a Fellow of
the Royal Society in recognition of his
distinction as a scientific navigator and
hydrographer: his sponsors included Charles
Darwin. Later he began the organization of
what became the British Meterological Office. It
is unfortunate that he should be remembered, if
at all, for his quarrel with Darwin and his tragic
death rather than for his considerable scientific
accomplishments. For more than a century his
meticulously drawn charts served scientists and
others navigating in the hazardous waters of the
Galapagos. R. D. Keynes** sums up FitzRoy's
varied achievement:

He deserves to be remembered not just as
Darwin's captain on the Beagle, although
the importance of the help and
encouragement that he gave during the
voyage, and his role in stimulating
Darwin's ideas, are not to be lightly
dismissed. He was also a hydrographer in
the front rank, parts of whose charts of
South American waters and sailing
directions for them are still in use nearly
150 years after the survey was conducted.
Above all he was one of the principal
founders of the science of meteorology.

-G. T. Corley Smith

* Reprinted with permission from Noticas de
Galapagos, 1985.

** The 6eag/e Record, edited by R. D. Keynes,
Cambridge University Press, 1979.

life, and so their collections were added to the
Galapagos catalog.

The Vanderbilt family enthusiastically
partook of this fashion. In the 1920s and 30s, the
collection of Galapagos species at the Centerport,
New York, Vanderbilt Museum grew, thanks to
such activities as the George Vanderbilt South
Pacific Expedition of 1937. Vincent Astor took his
yacht, the Nourmahal, to the archipelago in 1930.
His expedition featured the first census of the

flightless cormorants, penguins (see box page 67),
and fur seals. Templeton Crocker conducted his
own expeditions for the California Academy of
Sciences aboard the Zaca in 1932 and again in
1935. This comfortable yacht was equipped with
an aquarium with fresh seawater circulation, so that
live specimens could be brought back to the
Steinhart Aquarium in San Francisco, home of the
California Academy of Sciences.

A series of expeditions to the Galapagos

The Velero III. (from 1939 Hancock Pacific Expedition
Report)

organized by millionaire G. Allan Hancock began in
1927, aboard the Oaxaca. Hancock was particularly
interested in taking scientists and students from
different universities and museums on research
cruises. This series of voyages also featured the
very comfortable and specially equipped motor
yacht Velero III, and lasted until 1938.

The Galapagos held a special significance for
Hancock, and the results of the series of cruises are
described in the impressive volumes; "The Allan
Hancock Pacific Expeditions," published by the
University of Southern California Press in Los
Angeles. The littoral observations and collections
were valuable additions to the knowledge of the
islands' algae, hydrozoans, corals, flatworms,
threadworms, sea cucumbers, copepods, water
fleas, and crabs.

Even President F. D. Roosevelt figures into
the history of biological exploration in the
Galapagos. He made a long cruise through the
Pacific in 1938 aboard the U. S. S. Houston. This
expedition sampled some interesting marine fauna;
particularly sponges, soft corals, sea urchins, and
brittle stars.

New Techniques

With the availability of modern SCUBA techniques,
a whole new world opened up for marine research
expeditions. Animals could be collected selectively,
and behavioral studies could be made for the first
time. One of the first expeditions using SCUBA in
the Galapagos was the International Institute for
Submarine Research voyage of 1953-54. Among
the many later expeditions was that of the Italian
Gruppo Richerche Scentifiche e Tecniche
Subacquee. In 1973, the Italians made important
contributions to the knowledge of phytoplankton
biomasses and primary production in the
archipelago.

In 1976, the Scripps Institution of
Oceanography, using their Deep-Tow vehicle,
discovered thermal and chemical anomalies in the
Galapagos Rift Zone, north of the archipelago.
Subsequent, underwater photographs in 1977
disclosed dense benthic communities at 2,550
meters. These were dominated by previously
unknown species of large bivalves, tubeworms, and
crabs. The rift zone site was reviewed in 1977 (see
Oceanus, Vol. 20, No. 3) with the unmanned
vehicle Angus, and the deep-sea manned
submersible Alvin, operated by the Woods Hole
Oceanographic Institution (WHOI). This work led

to several universities and institutions organizing
the joint Galapagos Rift Biology Expedition in
January 1979 (see Oceanus, Vol. 22, No. 2). The
biological discoveries of this voyage were
supported by detailed descriptions of topography,
bathymetry, geology, and chemistry.

A turning point in Galapagos history was
reached with the Xarifa Expedition of 1953-54, led
by Hans Hass. He was a pioneer in discovering the
frontiers of SCUBA diving, and took photographs
and films of the underwater life. He and his
colleague Irenaus Eibl-Eibesfeldt, who came back
to the islands on his own in 1957, issued alarming
reports on the status of the endemic fauna. These
reports focused world attention on the Galapagos
problem," and led to the establishment of the
Charles Darwin Research Station, which began
operation in the early 1960s.

The Charles Darwin Research Station

The problems confronting the terrestrial fauna and
flora of the Galapagos in 1960 were immense. But
it was questionable whether there were similar
problems threatening marine species and habitats.
These questions were addressed during a
Galapagos symposium held as part of the 10th
Pacific Science Congress in Honolulu in 1961.

Another early assessment of the marine
environment was organized in 1964. The Charles
Darwin Foundation, the University of California,
and the California Academy of Sciences
participated in this cruise, the Galapagos
International Scientific Project. These studies
helped to set priorities for the first scientists at the
Research Station.

The first marine scientist at the Darwin
Station was Guy T. Houvenaghel in 1968. He wrote
his thesis in 1974 on the oceanography and
geomorphology of the Galapagos marine
environment. With his wife, Nadine, he described
plankton and shoreline zonation. The next marine
scientist was Gerard M. Wellington, now at the
University of Houston, Texas. He came to the
archipelago as a Peace Corps volunteer from 1973
to 1975. Wellington started an inventory of the
Galapagos coastal marine environments, and
recognized their vulnerability. He therefore
proposed the inclusion of marine areas within
existing national park boundaries. His findings and
recommendations included many suggestions for
the management and interpretation of a marine
park, and were presented to the Department of
National Parks and Wildlife in Quito.

Derek Green took over from Wellington in
1975 and continued a study on the ecology of the
East Pacific green turtle (Che/on/a mydas agassizi),
started by Peter Pritchard and Miguel Cifuentes.
The latter, who wrote his thesis on this project,
subsequently became Galapagos National Park
Service Superintendent from 1976 until 1986. This
turtle project was the first major program of the
station with the Ecuadorian National Fisheries
Institute (INP) in Guayaquil, with which a close
cooperation developed after 1976. Many
Ecuadorian students took part in the turtle project,
among whom was Mario Hurtado, who later

worked as Assistant Director at the station from
1984 to 1986.

Studies by IMP personnel were conducted
from 1976 to 1980 on the artisanal fishery of
bacalao, mullet, and three species of lobster — red
(Panulirus penicillatus), blue (P. grac/7/s), and slipper
(Scyllarides astori). The present Director of the
station, Gunther Reck, established and carried out
this project in the Galapagos for IMP. Plankton
samples have been collected extensively by both
IMP and the Oceanographic Institute of the
Ecuadorian Navy (INOCAR).

In March 1979, the station created the
Department of Marine Biology and Oceanography.
Two years later, Gary Robinson began work as
Marine Biologist. Until he left in 1983, Robinson
was mainly concerned with the planning and
building of the Marine Laboratory. Priscilla
Martinez assisted Robinson in a study of two
species of black coral; Antipathes panamensis, the
commercially exploited species, and its endemic
relative, A. galapagensis. Robinson also co-edited,
with Eugenia del Pino of the Catholic University of
Ecuador, the important volume of articles
published by the Charles Darwin Foundation, El
Nino in the Galapagos Islands: The 1982-83 Event
(see article page 42).

Non-station scientists have carried out
several long-term studies as well. Andrew Laurie's
marine iguana studies (see article page 54), begun
in 1980, are still continuing; although they will now
be taken over by Thomas Dellinger of the Max
Planck Institute in West Germany. From the same
institution, Fritz Trillmich has come to study fur
seals and sea lions since 1976, and continues to do
so.

The impact on the marine environment by
the declaration of the Marine Resource Reserve
(see decree page 4) remains to be seen. At the
moment, a management plan is being developed,
with personnel from various agencies in Ecuador,
and help from the United States and Australia.

The Future: Plans and Problems

The Darwin station has a budget of about $1
million, and is staffed by between 50 and 60
people. This puts tight constraints on the scope
and depth of the station's work. Therefore, the
station's upcoming studies will be focused on
providing the necessary information for decisions
about the management plan for the Marine
Resource Reserve. For example, assays of human
impact will include the exploitation of black coral,
artisanal fisheries, waste management, and tourist
boating activities. In cooperation with national
institutions, biological and chemical studies are
planned, or have begun, to assess nutrients, zoo-
and phytoplankton, petroleum concentrations, and
cetacean activities. An inventory will be made of
potentially threatened habitats, such as diving sites
and beaches.

Many problems face marine researchers in
the Galapagos. The station's marine laboratory, due
to limited financial possibilities, is only basically
equipped. Part of the diving gear has been donated
by, or bought from, departing scientists, so

The most exploited fish species in the archipelago is the
bacalao (Mycteroperca olfax), a large grouper, common
but endemic to the Galapagos.

equipment is not always compatible. Since there is
little available equipment on the Ecuadorian
mainland, it must be imported from the United
States or Europe. Thus, given the delays and
difficulties in obtaining materials and spare parts,
each study must be planned well in advance.

Nevertheless, present studies by visiting
scientists who use the equipment and
infrastructure of the marine laboratory have already
given some promising results. Peter VV. Glynn of
the University of Miami has published a book on
Galapagos corals and coral reefs with Wellington,
and is currently studying the recuperation of reef-
building corals after the devastation of El Nino. He
has indicated that predation, in particular by the
pencil-spined sea urchin, Eucidaris thouarsii, may
be limiting the recovery of some species. Although
Pavona and some others are recovering slowly,
Pocillopora is still not showing signs of a good
recovery.

In 1954, an area of Urvina Bay, on Isabela,
was uplifted very rapidly, exposing a shallow water
community. Mitchell Colgan of the University of
California at Santa Cruz, who is studying this area,
recently found a coral head with some 400 years of

Marine iguana (Amblyrhynchus cristatus).

The Charles Darwin Research Station

I he scientific interest in the Galapagos that
began with Charles Darwin continues today in his
name, through the work of the Charles Darwin
Research Station. The station — headquartered at
Puerto Ayora on Academy Bay, Santa Cruz
Island — is the offshoot and pride of the Charles
Darwin Foundation for the Galapagos Islands.
Since its beginning in 1960, the station has been
a valuable Galapagos resource for scientists and
tourists alike. But, had it not been for a few far-
sighted individuals 30 years ago, who
orchestrated public relations and diplomatic
activities around a set of coincidences, it might
never have happened.

The young Belgian ethologist (behaviorist),
Irenaus Eibl-Eibesfeldt, visited the Galapagos early
in the 1950s, as did San Francisco State University
ornithologist Robert I. Bowman — then a graduate
student. Both reported what they saw as the
impending doom of the stark beauty and
ecological integrity of the islands. The
unfavorable publicity about the cruel conditions
prevailing in the prison on Isabela Island
prompted the Ecuadorian government to close it
and avoid further political embarassment.
Another danger was the increasing numbers of
feral goats, pigs, dogs, and cats threatening the
irreplacable populations of endemic rodents,
finches, tortoises, and unique plants.

In 1957, the United Nations Educational,
Scientific, and Cultural Organization (UNESCO),
the New York Zoological Society, the
International Committee for Bird Protection, and
Time, Inc., sponsored a biological reconnaissance
of the Galapagos under the leadership of
Bowman and Eibl-Eibesfeldt. On their return, they
recommended a series of protective measures for
the islands' biota, and selected a site on Tortuga
Bay, Santa Cruz Island, for a future research
station — a site later abandoned in favor of the
one on Academy Bay.

According to Bowman, the centennial of
the publication of Darwin's the Origin of Species
in 1959 was the impetus to take the
recommendations of the 1957 expedition to
heart. UNESCO's first Director-General, lulian
Huxley, presided over a committee determined to
realize the ideas recently expressed, ideas that
first had been voiced in the 1 930s by California
Academy of Sciences ornithologist Harry S.
Swarth and National Academy of Sciences' Pacific
Science Board then chairman, Harold /. Coolidge.
The committee included Smithsonian Institution
Secretary, S. Dillon Ripley; Royal Academy of
Belgium Director of Science, Victor Van Straelen;
future President of the International Union for the
Conservation of Nature, Harold Coolidge;
founder of the World Wildlife Fund, Sir Peter
Scott; and Bowman, who served as Secretary-for-
the-Americas of the Charles Darwin Foundation

for the Galapagos Islands, which later that year
was incorporated under Belgian law. Veteran
biologist of the Belgian Congo Van Straelen
served as first president.

Bowman points to Coolidge especially as
the one who made it all happen. "If not for him
we might never have done it. He was a
remarkable diplomat, able to enlist the help of
prestigious organizations," such as the National
Academy of Sciences, National Science
Foundation, Royal Society of London, and the
Max-Planck Institute, "and make all the right
political connections."

The right political connections were a
necessity, if the various funding sources and the
Ecuadorean government were all going to act
smoothly together. Coolidge, with the help of the
first two secretaries-general of the CDF, lean
Dorst and lacques Laruelle, was able to pull the
right strings, and in 1959 Ecuador endorsed plans
of the CDF by declaring all uninhabited areas of
the archipelago a national park. Since there was
no national park service at the time, the
Ecuadoreans gave all administrative authority to
the Charles Darwin Research Station (CDRS),
which was established on Academy Bay by the
CDF in 1960.

The CDF chose a young Swiss
ornithologist, Raymond Leveque, to be the first
director of the CDRS. Bowman credits Leveque
with admirably facing the early challenges of the
station. He had to oversee housing and road
construction, choose the boundaries of reserve
zones, and specify which native plants and
animals required priority in conservation.

By 1 964 the early work was finished and
the CDRS was officially inaugurated. To insure
that all the funding organizations and political
bodies involved would enthusiastically support
the station, according to Bowman, the CDF
planned for the grandest celebration the
Galapagos may have ever seen. The United States
Navy, Air Force, and Army all got involved.
Distinguished scientists from England, japan,
Belgium, the United States, and elsewhere, and
other notables such as the artist Roger Tory
Peterson, and the grandson of Charles Darwin,
Commodore Barlow, were on hand for the
dedication ceremonies at Academy Bay, Santa
Cruz Island. The Ecuadorian government was
represented by various ministers, including
General Marcos Gandara, a leading member of
the then ruling Military junta of Ecuador. The
general, a former engineering professor at the
Catholic University in Quito took a special
interest in the activities of the scientists, often
seen riding the shells of the giant tortoises housed
in display pens at the Darwin Station. Largely
through the personal attention of General
Gandara, an executive decree was issued

establishing special reserves for the protection of
Galapagos wildlife. So committed was the
government of Ecuador to the concept of
inviolate nature reserves in the Galapagos that
they stood by their decision, in the face of strong
opposition, to relocate settlers illegally farming in
protected regions of Santa Cruz Island. "Of
course," says Bowman, "after all this, there was
no way anyone could back down, and we were
off and running."

Because of the enthusiastic support of the
Ecuadorean government, the station was able to
move quickly on important conservation issues,
such as feral animal control and surveillance of
private yachts. The close ties of 20 years ago have
been nurtured by the station, and many
Ecuadoreans have served on the station's staff.

Effective cooperation between the CDRS
and Ecuador has been a big factor in the success
of both the station and the Galapagos National
Park Service (GNPS), founded in 1 968 — again
according to Bowman. In the late 1960s, CDRS
director Roger Perry developed a program that
brought Ecuadorian students, local school
teachers, tourist guides, and scientists for
seminars on conservation and research projects in
the Galapagos. Many of these later became
crusaders for conservation in continental Ecuador
and elsewhere in Latin America. The educational
program of the Darwin Foundation was
successful. The cause of wildlife protection in
Ecuador had a large endemic following.

Because so many young Ecuadoreans have
had the opportunity to learn the methods of
conservation and science in these remarkable
islands, under the microscope of the global
conservation community, there is a growing
belief that Ecuador now has one of the finest
national parks in the world. "They've been well-
trained, and now they're great political fighters,"
Bowman says. "If we ran Yosemite as well as they
do the Galapagos, we'd be better off." Other
educational activities of the station include
training GNPS guides and wardens, and natural
history instruction for local teachers.

In the area of conservation, the efforts of
the CDRS have been most successful. No known
extinctions have occurred on the islands since the
1950s, although certain island populations of
widespread forms have died out. For the most
part, the feral animal situation is better
controlled. In 1 965, responding to the
decimation of tortoises in certain areas, the
station started a captive breeding colony.
Beginning with only two males and ten females,
they have now reintroduced more than 100
tortoises to Espahola Island.

Because of the heavy administrative load
on the CDRS staff, most scientific research in the
Galapagos is carried out by visiting scientists. The
station is intimately involved with all the ongoing
research, however. Through regulating the areas
where research is conducted, and giving logistic
and technical support to visiting researchers, the
scientific interest of the Galapagos has remained
high — attracting a growing number of scientists
of many disciplines.

Presently the Darwin station employs a
staff of 80 people, and about 50 visiting scientists
work at the station each year — most work for just
a month or two, but some have stayed for up to 2
years. Facilities of the station now include the 70-
foot research vessel Beagle III, a reasonably well-
equipped laboratory, darkroom, workshop,
meteorological and seismographic station, and
living accommodations. For complete details,
scientists interested in working at the station or
obtaining visiting scientist status should contact
either

Gunther Reck, Director Juan Black, Secretary-
Charles Darwin General
Research Station Charles Darwin
Casilla 58-39, Foundation
Guayaquil, Ecuador Edificio Colon

Avenue Colon 535-y 6
de deciembre
Quito, Ecuador

— TMH

continuous growth. Study of this area may permit
accurate reconstruction of past El Nino events (see
article page 61). This year Hal Whitehead, of
Dalhousie University in Halifax, Canada, is
continuing his studies on the social behavior of
sperm whales. His 1985 survey gave very
interesting and promising results. So the work is
continuing (see article page 49).

The conservation aspect of studies done by
the station is a driving force far greater than pure
scientific curiosity. Nonetheless, the work of the
station has demonstrated that a successful
combination of the two can be found. The
Galapagos underwater ecosystem is unique in its

combination of high species complexity, a wide
diversity of species, and a high degree of
endemism. The situation of the archipelago on a
junction of major current systems, and its wide
diversity of habitats, serve to create a distinct
regionalism of the islands. Hopefully, the Marine
Resources Reserve can protect these features, and
give a new impulse to marine research in the
waters of the Galapagos.

Henk W. Kasteleijn is head of the Department of Marine
Ecology and Oceanography of the Charles Darwin
Research Station, Santa Cruz, the Galapagos, Ecuador.

Negative Effects

of the 1982-83 El Nino

on Galapagos Marine Life

by Gary R. Robinson

Dy March 1983, it was evident that the world's
weather was in the grip of the strongest El Nino/
Southern Oscillation (ENSO) event ever recorded.
El Nino conditions in the eastern Pacific were
associated with droughts in Australia and Africa,
cyclones in French Polynesia, and intense storms
along the Pacific coast of North America. With
almost daily exposure in newspapers and
periodicals, El Nino became a household word.

The isolated Galapagos Islands are the only
emergent land forms in the eastern Pacific Ocean
lying directly in the path of developing El Ninos. Not
surprisingly, the 1982-83 event had profound effects
on the archipelago's biota. A nutrient-poor blanket
of warm tropical water invaded the islands,
depressing the normally shallow thermocline* to
great depths. Dissolved nutrients, such as nitrate,
phosphate, and ammonia, usually found below the
thermocline, were removed from the range of
upwelling waters around the islands. Reduced
availability of nutrients in the sun-lit surface layers
led to reduced primary productivity, which
reverberated through all levels of the trophic food
web.

The effect of declining and shifting food
resources was most pronounced at higher trophic
levels, especially for seabirds, pinnipeds — sea lions
and fur seals — and marine iguanas. Each year these
animals provide nearly 26,000 visitors per year to the
Galapagos National Park with some of their most
memorable experiences.

Tourism Hit Hard

The strength of the 1982-83 El Nino event was
dramatized by a request made by a tour company to
myself and other scientists at the Charles Darwin
Research Station located on Santa Cruz Island. The
request was for a document describing what was
occurring to the populations of marine iguanas, sea
lions, fur seals, and seabirds that visitors were

* A zone where the water temperature decreases more
rapidly than the water above or below it. This zone usually
starts from 10 to 500 meters below the surface and can
extend to over 1,500 meters in depth.

accustomed to seeing. How long might El Nino last,
and when would the animal populations recover?
The document was to be distributed to travel agents.
The 1 982-83 El Nino hit tourism hard. News of the
conditions in the islands traveled fast and far.

Stories of marine iguanas dying at visitors'
feet, of fewer or no seabirds along park trails, and
starving fur seals circulated in the travel business.
The problem was, the stories were true.
Consequently, fewer people decided to travel to the
Galapagos in 1983, preferring instead to delay their
plans for some other year when conditions might be
closer to normal.

The document requested by the tour
company was never prepared. We had no idea
when El Nino conditions would ameliorate, nor did
the many scientists then working in the islands know
precisely how the native fauna and flora might
respond to this unprecedented event. There had
been El Ninos before; the last in 1976 — but none on
the scale of the 1982-83 event. Change was evident
everywhere in the islands, and the excitement of
simply observing and recording the results of it
infused everyone's conversations. El Nino and its
associated conditions were providing clues of how
mechanisms of natural selection might operate,
which, over the course of a few million years, had
led to the intriguing flora and fauna inhabiting these
islands today.

El Nino Sets Records

In terms of weather records, the 1982-83 event
exceeded all previous marks. Climatological records
at Academy Bay on Santa Cruz Island have been
collected since 1965 by the Charles Darwin
Research Station and furnish the basis for
comparison.

The sea-surface temperature (SST) anomaly
rose to 4 degrees Celsius above normal by
December 1982 and stayed at that level until May
1983, when the anomaly rose to +4.5 degrees
Celsius. By June, it reached a record +5.5 (Figure 1).
SSTs then quickly dropped, approaching normal
values by September 1983, when El Nino subsided.

Record rainfall fell on the Galapagos
coincident with the warm tropical waters bathing the

2 2

1 0-

,, 28-

26-

2 24-
a

.* 22-

JJASONDJ
1982

MAMJJASOND
1983

Figure 1. (A) Sea-surface temperature anomaly measured near
Academy Bay, Santa Cruz Island, during the 1982-83 El
Nino. Maximum anomaly exceeded +5 degrees Celsius in
June 1983. Dotted line at zero indicates normal sea-surface
temperature. (B) Monthly mean sea-surface temperature
during the 1 982-83 El Nino compared to the 1 7-year mean
(1965-1981) at Academy Bay. (After Kogelschatz, and others,
1985)

islands. Between the months of November 1982 and
July 1983 (9 months), more than 3,225 millimeters
(126.9 inches) of rain fell at the coastal Santa Cruz
Island weather station (Figure 2). Annual rainfall
averages 374 millimeters (14.8 inches), so El Nino
dumped more than nine times the usual amount of
precipation. The heaviest recorded rain during the
event was 136.7 millimeters (5.4 inches) on
December 17. The normally arid Galapagos Islands
were transformed into lush tropical islands, with an
exuberant growth of entangling vines. Roads became
quagmires of mud, and were often interrupted by
rushing rivers after torrential rains. The floor of
several volcanic craters became giant cisterns of
fresh water (Figure 3).

Both the record rainfall and the rise in sea

E
E

600-

400-

DC 200-

JFMAMJJASONDJFMAMJ JASOND
1982 1983

Figure 2. Rainfall recorded at the Charles Darwin Research
Station on Santa Cruz Island for the 1982-83 El Nino. Total
rainfall between November 1 982 and July 1 983 measured
3,225 millimeters (126.5 inches). Solid line with monthly dots
is average rainfall for the period 1979-81, bars are actual
rainfall in 1982-83. (After Robalino, 1985)

Figure 3. North crater of Pinzon Island during a normal year
(A) and filled with water during the 1982-83 El Nino (B).
(Photos by Linda Cayot)

level associated with El Nino were bimodal, with
minimum values during February 1983 (compare
Figures 2 and 4). The first pulse in rising sea level was
attributed to the relaxation of sea-surface slope
across the Pacific basin due to reduced tradewinds.
A second pulse followed when anomalous westerly
winds blew over a large portion of the central Pacific
from January to May. Surface waters moved
eastward toward the Galapagos, accounting for the
second peak in sea-level rise around May 1983.
These winds also sparked the unusual cyclones that
hit French Polynesia in 1983.

Positive sea-level anomalies in the Galapagos
indicated a thickening of the warm layer and
increased heat content of the ocean surrounding the
islands. Warm air saturated with moisture from the
ocean's surface released precipitation as these air
masses were forced upward by the islands. Several
storms during 1983 were especially severe. Lightning
had never before been observed by long-term
residents of the Galapagos (a period of 45+ years).
But, in 1983, lightning bolts felled palm trees and
wreaked havoc on municipal power stations.

The marine environments of the Galapagos
are poorly understood, as is the biology of most of its
marine organisms. Numerous observations
throughout the Galapagos during El Nino indicated
wide perturbations in abundance, and altered

JFMAMJJ ASOND|JFMAMJ JASONDJFMAMJJ ASOND

1981 1982 1983

Figure 4. Daily mean sea-level
anomaly at Santa Cruz from
1981 to 1983. (After Wyrtki,
1985)

patterns of distribution of marine species among the
islands. Since normal year-to-year variation in
population size is largely unknown for many
organisms, it is difficult to judge the impact of the
1982-83 El Nino. However, our knowledge of
seabirds, marine iguanas, pinnipeds, and coral
communities is more complete, as the following
examples show.

Waved Albatross

Apart from a few errant pairs, the waved albatross,
Diomedea irrorata, breeds only on the tiny
Galapagos Island of Espanola, in the southern region
of the archipelago. Catherine Rechten, then
associated with the Max Planck Institute in Germany,
returned there in March 1983 to find torrential rains,
a thick cloak of vegetation over usual nesting areas,
and diminished food resources. She blamed the
circumstances for the complete breeding failure of
the waved albatross in 1983.

Several observations indicated that the waved
albatross experienced food shortages. Adult birds
arrived at nesting sites approximately two weeks
later than usual. Far fewer adults laid eggs in 1983
than in 1982, and the size of eggs was noticeably
smaller.

In a normal year, roughly a third of the nesting
females are less than 10 years old. But in 1983, all of
the egg laying birds at the study site were older than
10 years and none of them had raised a chick to
fledging the previous year. In other words, raising a
chick demanded considerable expenditures of
energy for parent birds, which could not be fully
recovered between breeding seasons because
El Nino intervened. Some older individuals, having
failed to fledge a chick the previous year and facing
the end of their reproductive life, could not pass up
the opportunity at breeding — even under the
marginal El Nino conditions. Younger birds could
afford to forego one laying season as many
reproductive years still lay ahead.

Unfortunately for the waved albatrosses
attempting to breed in 1983, El Nino soon made it
impossible. Not one egg hatched, and all eggs laid in
1983 were abandoned within 20 days. Eggs normally
hatch in about 63 days.

Rechten describes adult birds swimming on

the surface of pools, trying in vain to retrieve eggs
after heavy rains submerged them. But rains were
not the only factor contributing to egg
abandonment. Male birds returned much later from
their foraging period at sea to relieve partners from
egg incubating stints; and, when they did return, it
was already too late.

For visitors to the islands though, the breeding
failure of the waved albatross had desirable
consequences. As Rechten writes, "Compared with
other years, courtship dances were unusually
frequent, as all adult pairs had failed to breed and
were therefore dancing."

Birds were not celebrating release from
parental duties. Rather, the courtship dance and bill
fencing that the waved albatross is so noted for
serves as a behavior that strenthens the bond
between mates (see box page 67). Waved
albatrosses typically engage in this behavior at the
end of the breeding season, or after failing to raise a
chick and flying away to the open sea.

Did the 1982-83 El Nino severely affect the
waved albatross population? Apparently not. Based
on tagging data, a 1 7 percent success at fledging
chicks would be sufficient to balance normal
mortality. Rechten's estimates of breeding success
for the waved albatross were 46 percent in 1981, 8
percent in 1982, and 0 percent in 1983. Being a
long-lived bird, however, one good reproductive
year can make up for disastrous years, such as 1983.

Galapagos Pinnipeds

While waved albatrosses failed to produce any
young during the 1983 season, and may have
suffered slightly higher adult mortality because of
El Nino conditions, the population was expected to
rebound once conditions normalized. Galapagos
pinnipeds, on the other hand, were drastically
affected by El Nino, and the effect carried over to
the following reproductive season.

The Galapagos supports two species of
pinnipeds — the Galapagos sea lion, Zalophus
californianus wollenbaeki, a smaller subspecies of the
California sea lion; and the endemic Galapagos fur
seal, Arctocephalus galapagoensis. Fritz Trillmich, a
biologist with the Max Planck Institute, has had a
long interest in pinniped biology in upwelling

tropical ecosystems. For more than a decade, he has
been involved with studies of population dynamics,
diving behavior, and maternal-pup behavior of
Galapagos sea lions and fur seals. In the journal
Oecologia (1985), Trillmich and Dominique
Limberger described the 1982-83 El Nino's impact
on a population of fur seals on Fernandina Island:

El Nino decimated the population of fur seals by
almost entirely eliminating the four youngest year
classes (1980 through 1983). Adult females and
non-territorial males suffered about 30 percent
mortality. Large male fur seals, holding breeding
territories in the early months (August to
November) of El Nino apparently could not
recover weight lost in territorial defense as El
Nino conditions developed to their full extent
following the breeding season. Nearly 100
percent mortality was observed for these
individuals, which were replaced by smaller
males the following reproductive season.

How did El Nino exert its impact upon
Galapagos fur seals? Shortage of food seems to be
the answer, although the evidence is indirect. The
fur seals' preferred prey of squid and fish apparently
remained in the cool waters below the thick blanket
of warm tropical water, beyond their diving abilities
(greater than 50 meters). Unfortunately, no direct
sampling of the distribution of these prey has been
undertaken in the islands.

It was clearly evident that there was a food
shortage. Trillmich and Limberger attribute the large
number of stillbirths in 1982 to nutritive stress.
Females that did pup successfully, stayed longer at
sea between nursing periods, and pups were
undernourished. In fact, none survived El Nino. The
year classes of 1980 and 1981 also suffered high
mortality. Young fur seals are dependent on their
mothers for nourishment for the first two years of
life, and do not gain full independence until a year
or two later.

In effect, the strong El Nino of 1982-83
produced a big hole in the age structure of the fur
seal population that may have large implications for
the future reproductive output of this species. In the
1983 reproductive season following El Nino,
surviving adult females had still not gained sufficient
resources to support added reproductive costs as
evidenced by low birth rate (1 1 percent of normal)
and significantly lower pup birth weights.

The effect of El Nino on the Galapagos sea
lions was similar to that of fur seals. Sea lion colonies
on the islands appeared deserted in 1983 compared
to former years, and most of the 1982 year class
died, apparently abandoned by adults remaining at
sea longer in search of diminishing resources.
Compared to fur seals, food shortages may not have
been as drastic for sea lions, as this species is
capable of deeper diving. But, like fur seals, pup
production in the year following the 1982-83 El
Nino was much lower for the Galapagos sea lion.

Low food availability may not have been the
only reason for the decline in sea lion populations.
Nutritive stress and physiological stress due to the

Figure 5. Young Galapagos sea lion with pox sores.
Prevalence of this disease was greater during El Nino due to
stress caused by elevated air and sea temperatures and by the
scarcity of food. (Photo by Gary Robinson/VU)

warm ocean and atmosphere may have made
individuals more susceptible to disease, particularly
to sea lion pox (Figure 5). The prevalence of this
disease seemed to be greater in 1983 than in
previous years and may have been a contributing
mortality factor.

Marine Iguanas

No creature epitomizes the uniqueness of the
islands more than the marine iguana, Amblyrhynchus
cristatus (Figure 6). When Charles Darwin visited the
Galapagos islands in 1835, he described the iguana
as "a hideous looking creature, of a dirty black
colour, stupid, and sluggish in its movements."

In November 1982, Andrew Laurie (see article
page 54) of Cambridge University, England, began
his third year of observations of a large colony of
marine iguanas on Santa Fe Island. At this site,
marine iguanas are largely intertidal feeders, moving
out onto exposed rocks as the tide drops to feed on
leafy green and red algae. With increased sea
temperatures and low nutrients because of El Nino,
most kinds of algae were replaced by filamentous
brown algae (principally Ciffordia sp.) that colonized
rocky substrates throughout the islands.

Marine iguanas continued to feed on the
brown algae, but rapidly lost weight. As El Nino
conditions persisted, marine iguanas starved to
death, even though Laurie's examinations of dead
animals revealed guts packed with filamentous
brown algae. Subsequent analyses of the
digestability of Ciffordia, compared to red algae
normally available to iguanas, indicated that the
brown algae offered far fewer metabolizable calories.

Figure 6. Marine iguana populations throughout the islands
were reduced 30 to 55 percent under El Nino conditions.
Usual species of algae consumed by iguanas disappeared
during 1982-83 and were replaced by a relatively
unnutritious species of filamentous brown algae.

In anthropomorphic terms, it was as if El Nino had
replaced a meal of steak, potato, and vegetable with
soda crackers.

The marine iguana population at Santa Fe was
reduced by about 65 percent. The 1983 hatchlings
were relatively unaffected by El Nino conditions,
probably because yolk sac reserves carried them
through the adverse conditions. In contrast, the 1982
year class suffered nearly 90 percent mortality since
small iguanas are entirely dependent on intertidal
feeding. Increased sea level and heavy swells
associated with El Nino denied them access to usual
feeding areas, which, in any case, were depleted of
algae.

An island-wide survey made following El Nino
indicated reductions of 45 to 70 percent in marine
iguana populations. How severely populations on
different islands were affected depended on the
extent to which iguanas fed subtidally at different
sites. Subtidal diving iguanas showed higher
survivorship, presumably because nutritious species
of algae were still available.

El Nino's influence carried over into the
following reproductive season. Normally, more than
1,800 females dig nests at the Santa Fe study site,
but only 10 did so during the nesting season

following El Nino. Allowing for 50 percent mortality
of adult females, the 10 nesting females represented
only 1 .2 percent of possible egg laying females.
Surviving female marine iguanas were not able to
acquire sufficient resources to compensate for the
30 percent weight loss incurred during El Nino in
time for the following nesting season.

Hermatypic corals

One of the more pervasive impacts of the 1982-83
El Nino was the widespread bleaching and mortality
of hermatypic (reef-building) corals throughout the
eastern Pacific. Peter Glynn, then with the
Smithsonian Tropical Research Institute in Panama,
and now with the University of Miami, was the first
to report on this phenomenon occurring at coral
reefs in the Gulf of Chiriqui, Panama. At the same
time as his observations, corals began bleaching
(losing pigment and symbiotic algae) in the
Galapagos during February 1983. By the end of
March, visitors to all sectors of the Galapagos were
reporting "dead" corals. As El Nino progressed,
bleached corals did indeed begin to die and coral
skeletons were overgrown with filamentous algae
(Figure 7).

Galapagos corals appeared to be very
sensitive to the anomalous oceanographic conditions
and exhibited very high rates of mortality. Alabaster
coral, Pocillopora sp., was most affected. In areas
where this coral forms incipient reef framework,
such as Onslow reef, all coral heads died and
became carpeted with filamentous algae.

The massive corals Pavona clavus and Por/tes
lobata also exhibited high levels of mortality. The
fact that massive coral colonies measuring more than
1 meter in height died during El Nino, coupled with
their known growth rates of 6 to 12 millimeters per
year suggest just how unusual the event was. Such
corals were easily three quarters of a century old.

Why the corals died is a different matter,
though. In general, corals are expected to fare better
under El Nino conditions. Clearer and warmer waters
should permit algal symbionts residing in the coral's
tissue to photosynthesize at higher rates. Corals are
able to utilize some of the products produced by
their symbionts, and consequently should grow
faster. But the positive relationship of increased
temperature to increased coral growth rate only
exists up to a certain point. Above 29 degrees
Celsius, coral growth rate rapidly declines. Factors
such as increased light intensities and reduced
salinities exacerbate problems corals face under
persistent elevated temperatures. All of these factors
were present in the Galapagos during the 1982-83 El
Nino and contributed to the extensive mortality
observed there.

Despite its equatorial location, coral
development is not very extensive in the Galapagos
Islands because of the relatively cool waters that
normally encircle the islands. Marine ecologists Peter
Glynn and Gerard Wellington noted, however, that
incipient coral reef formations and corals themselves
are surprisingly common in the Galapagos marine
environment. The prevalence of corals in the islands
was made visible during El Nino because bleached

Figure 7 . Bleached coral head
of Pavona clavus near
Academy Bay, Santa Cruz
Island during El Nino. Corals
bleached (lost pigment and
symbiotic algae) by February
1983, but were not colonized
by filamentous algae until April
1983. (Photo by C. Robinson/
VU)

heads contrasted markedly against the background.
It is ironic that Clynn and Wellington's detailed
descriptions of Galapagos coral reefs was published
in the same year El Nino was decimating such
formations.

These events suggest one reason why coral
reefs in the Galapagos may remain "incipient."
Perhaps every century or two, strong El Nino events
occur causing extensive coral mortality. Physical and
bioerosion of dead coral framework, particularly by
the pencil urchin, Eucidaris thourasii, may reduce the
structure to rubble before living corals are able to re-
establish and begin the reef building process again.

A Positive Footnote

El Nino/Southern Oscillation events are generally
associated with negative impacts and the examples
related here do not dispel that notion. For many
marine species dependent on productive Galapagos
upwelling ecosystems, El Nino had adverse impacts.
Examples are the penguins, flightless cormorants,
pinnipeds, marine iguanas, and blue-footed boobies.
But for species with tropical affinities, such as
moorish idols, Zanc/us canescens, and filefish, Aleutra
inschpta, the warm water conditions eliminated
temperature barriers, and these species dispersed
throughout the islands.

Rarely occurring El Nino events of the
intensity of the 1 982-83 phenomenon may set the
stage for the production of new species. They push
populations to the brink of extinction under their
adverse conditions. But once these conditions abate,
small founding populations may remain. Isolation,
mutation, and genetic drift may operate on these
small founding populations, making possible the
evolution of new species.

In another respect, adults or dispersal stages
of marine species may accidentally come to the
Galapagos because of the unique oceanographic
setting — strong easterly transport of surface waters
from the Indo-Pacific region, and more commonly,
southerly transport via the Panama Current during
El Nino conditions. The prolonged conditions of the

Galapagos Crossroads

\-Jceanographic and climatic variability are
hallmarks of the Galapagos Islands. The islands lie
along the equator about 1,000 kilometers west of
mainland Ecuador in the transitional zone of the
eastern tropical Pacific Ocean. Several major
ocean currents run side by side here like east-
and west-bound lanes of a major freeway, with
the Galapagos Islands representing a crossroads
intersection (see map on following page).

These current systems vary in intensity
during the year in a seasonal see-saw which gives
two distinct seasons to the Galapagos. They are a
wet season, extending from January to March
with rainfall amounts of 200 to 370 millimeters at
coastal stations; and a dry season, during the
months of June to December, characterized by
strong local winds, cooler sea temperatures, and
a persistent high fog known locally as garua
(Spanish for mist).

During the dry season, the South
Equatorial Current (SEC) is well-developed and
brings cool water (18 to 22 degrees Celsius) to
the islands. This current is derived from the Peru
Oceanic Current and Peru Coastal Current, which
flow along the west coast of South America. As
these water masses approach the Equator, they
turn westward to become the SEC. The flow of
these currents is determined mainly by the
strength of the southeast tradewinds, which
generally blow strongest during the months of
August and September. The SEC sweeps along
the entire width of the Pacific, beginning just east
of the Galapagos and ending in the western
Pacific. As the SEC wafer mass is transported
across the Pacific, it is gradually heated under the
equatorial sun, setting up a sea-surface
temperature (557) gradient of 4 to 5 degrees
Celsius from west to east. The gradient in 557 is

North Equatorial Countercurrenl

Panama
Current / /

Galapagos Is

South Equatorial Currem *•'•

Equatorial
Undercurrent

Peru Oceanic Current
Peru Coastal Current

Pacific Ocean

also reflected in the depth of the thermocline,
which may be deeper than 125 meters in the
western Pacific and near the surface in the
Galapagos region. Jradewinds that blow across
the Pacific, driving the SEC, also set-up and
maintain a basinwide tilt to the ocean's surface.

The Equatorial Undercurrent (EUC, also
known as the Cromwell Current) is prevalent in
the Galapagos during the dry season. This current
flows beneath the SEC, but in the opposite
direction, from west to east. As the EUC
approaches the Galapagos platform it is deflected
upwards, bringing cool nutrient-rich waters to the
surface.

The intense upwelling and highly
productive waters of the western region of the
Galapagos are distinctive of the EUC. Whales and
dolphins are common in this area, as are the
endemic flightless cormorant and Galapagos
penguin (see box page 67). Under the EUC
influence, sea-surface temperatures around the
northwestern tip of tsabela Island may measure
only 16 to 18 degrees Celsius — at the equator!

During the wet season, the SEC and EUC
slacken because of the weakening southeast
tradewinds, and the balance tips toward the
Northern Ecuatorial Counter Current (NECC)

Major ocean currents in
relation to the
Galapagos archipelago.
These currents vary
during the year,
producing a dry season
and wet season. The
Peru Current is also
known as the Humboldt
Current, and the
Equatorial Undercurrent
is also known as the
Cromwell Current.
(After Grove 1 984)

system. The Equatorial Front, the dividing line
between the NECC and the SEC, lies at
approximately 4 degrees North of the equator
during the dry season, and moves toward the
islands during the months of January through
March. Warm (25 to 28 degrees Celsius) tropical
surface waters of reduced salinity and nutrients
may flow out of the Panama bight as the Panama
Current. During this season, the winds are
generally mild, the seas calm, and the skies clear
except for the large cumulus clouds that
occasionally produce tropical downpours.

The convergence of three distinct water
masses at the Galapagos has transported
representative marine biota from tropical and
subtropical regions of Central and South America
as well as from the Indo-Pacific. The level of
endemism is quite high, averaging about 25
percent. Coupled with the broad range of habitat
types available (mangroves, precipitous cliffs,
sandy beaches, and so on) and the complex
interplay of currents dividing the various islands
into distinct zones, it is not surprising that a great
diversity of marine life resides in the Galapagos
seas. The Galapagos marine environment is a
"melting pot" of species that biogeographers
recognize as a distinct biotic province. — GR

1982-83 El Nino may favor this sort of recruitment,
and the successful establishment of tropical species.
These species may ultimately become permanent
members of the Galapagos marine communities,
adding to their diversity.

Gary R. Robinson is Manager of the Sea Center at the Santa
Barbara Museum of Natural History, and was formerly
Resident Marine Biologist at the Charles Darwin Research
Station in the Galapagos.

Acknowledgments

I wish to acknowledge the Charles Darwin Research
Station, Galapagos National Park, and the National Institute
for Galapagos for their support of marine research.
Contribution No. 415 of the Charles Darwin Foundation.

Selected References

Clynn, P. W., C. M. Wellington, and C. Birkeland. 1979. Coral reef

growth in the Galapagos: Limitation by sea urchins. Science 203:

47-49.
Clynn, P. W., and C. M. Wellington. 1983. Corals and Coral Reefs of

the Galapagos Islands with an Annotated List of the Scleractinian

Corals of the Galapagos. 297 pp. Berkeley: University of

California Press.

Grove, ). 1984. At the heart of El Nino. Oceans 17: 3-8.
Robinson, G., and E. M. del Pino, eds. 1985. El Nino in the

Galapagos Islands: The 1982-1983 Event. 533 pp. Contribution

No. 388 of the Charles Darwin Foundation for the Galapagos

Islands. Quito, Ecuador.
Trillmich, F., and D. Limberger. 1985. Drastic effects of El Nino on

Galapagos Pinnipeds. Oecologia 67: 19-22.
Wellington, G. M. 1984. Marine environment and protection. In Key

Environments Series: Galapagos Islands, eds. J. E. Treherne and R.

Perry. Oxford, England: Pergammon Press.

Sperm Whale Behavior

the Galapagos Grounds

In early 1985, we sailed our 10-meter sailboat/
research vessel to the Galapagos Islands to study
the social behavior of the sperm whale (Physeter
catodon). Most biologists who visit the Galapagos
are interested in the unusual, often endemic,
plants, animals, and biological systems that have
evolved there, or the pests that man has
introduced to the islands. But the sperm whale is
found throughout the world's oceans — it is one of
the most widespread mammals apart from man, his
pets, and other followers — and poses no known
threat to endemic Galapagos organisms. Then why
sail to the Galapagos to study sperm whales?

We were looking for a place where we
could follow the groups of female sperm whales—
the primary units of sperm whale society. These
"nursery" groups are generally restricted to tropical
and subtropical waters. We were hoping to
examine the interactions between these groups
and mature male sperm whales.

The large male sperm whales, which may

sometimes reach 18 meters in length and about 60
tons in weight — about three times the mass of a
mature female — are thought to spend part of the
year in cold, sometimes polar, waters, but
presumably return to the tropics to mate. However,
the details of this mating system are unknown, and
a possible source of error in the models of sperm
whale population dynamics developed by the
Scientific Committee of the International Whaling
Commission (IWC). Their "Sperm Whale Model"
assumes a harem system, with a large male taking
over a group of females and defending them
against other males. However, there is no concrete
evidence that such a system exists, and sometimes
more than one male has been found with a group
of females. This led several scientists to speculate
that male sperm whales might form long-term

Above, the head of a female sperm whale photographed
underwater. (Photo by the author)

coalitions with other males, and take over groups
cooperatively, in the manner of male lions
(Panthera leo).

Other questions arose when the South
African scientist Peter Best, based on a study of
parasites on sperm whale carcasses, suggested that
associations between breeding male sperm whales
and particular groups of females might be much
briefer than an entire breeding season. To give a
fair description of the social organization of sperm
whales and to construct reasonably accurate
models of their population dynamics, detailed
studies of free-living sperm whales are needed.

The "Galapagos Grounds"

Between 1982 and 1984, Jonathan Gordon of
Cambridge University in England and I had
developed methods of tracking sperm whales using
passive acoustics; of identifying, measuring and
sexing animals photographically; and studying their
diet from fecal samples. We studied sperm whales
in both the Atlantic and Indian Oceans, but found
no ideal area. We needed a study area with both
groups of female sperm whales and large males,
with calm water, and as few logistic and
bureaucratic hurdles as possible. After examining
whalers' records, survey reports, climatic charts,
and government regulations, with the help of my
colleagues, I decided to try the Galapagos.

The Galapagos Islands were first noted as a
sperm whaling ground by British Royal Navy survey
ships in the late 18th century. But it was the
Yankee whalers who developed the industry. In the
first half of the 19th century, whalers from
Nantucket, New Bedford, and other New England
ports took thousands of sperm whales from
Galapagos waters. The sperm whales were not the
only animals affected by these whaling operations:
Galapagos fur seals were slaughtered for their pelts
and oil, and large numbers of the giant Galapagos
tortoises were captured for food (see page 86). By
the 1860s, the Yankees found the Galapagos "dry
cruising" — presumably most of the sperm whales
had been taken. The Galapagos escaped the ultra-
efficient mechanized whaling of the 20th century,
and, apart from a few brief ship surveys, there was
little recent information about the sperm whale
population when we arrived in 1985.

We found the sperm whales principally in
the productive waters west and southwest of
Isabela (see map page 2), the largest island in the
archipelago, where the Cromwell Current, running
eastwards beneath the Equator across the Pacific,
finally meets land and is forced to the surface.
Captain Colnett, who headed one of the first
British exploratory whaling expeditions, had
recommended the same area "to all cruizers" in the
1790s.

Sperm Whale Groups

During our 2'/2 months in Galapagos waters, we
photographed slightly more than 200 individual
female and immature sperm whales. Using
statistical criteria, we clustered these into 13
recognizable groups, which appeared to have a
reasonably closed membership (no obvious

immigration or emigration) during the time of our
study. Most of these groups contained about 20
females and immatures and 1 to 2 small (3 to 4
meters long) first-year calves. The statistical analysis
also suggested that we encountered most of the
groups present in the waters off the Galapagos. We
estimated a total population of about 270 sperm
whales in the area at that time.

We were able to sail continuously with
groups of whales for up to 10 days. Occasionally,
as groups encountered one another, we sometimes
found ourselves leaving one group to follow a
second group that was known to us from a
previous encounter.

Two Modes of Behavior

The sperm whales appeared to have two principal
modes of behavior: "feeding" and "social." Most
frequently they seemed to be feeding. Each
member of the group would dive for approximately
40 minutes, and then surface to breathe for about
10. Traces of the whales' dives when seen on a
recording depth sounder showed them to be
diving almost always to 410 meters, ± 15 meters,
both day and night. Their choice of 410 meters is a
mystery at present. We have not yet identified any
obvious oceanographic features 410 meters below
the surface off the Galapagos. In other ocean areas,
sperm whales have been tracked diving to a variety
of depths. Analysis of fecal samples suggested that
the Galapagos sperm whales were generally eating
deep sea squids, so perhaps 410 meters has some
special significance for these prey.

While apparently feeding, a group would
usually spread out over several square kilometers
of ocean, often aligned in a rank perpendicular to
their direction of travel, apparently sweeping the
ocean for food. The whales would usually appear
at the surface for their breathing periods singly or
in pairs; but if other whales were within a few
hundred meters, they would often alter course to
join up. After their breathing period, together or
separately from their companions, the sperm
whales would often lift their flukes into the air to
begin the dive.

While at depth, a sperm whale usually made
a series of clicks, about one every 0.5 to 1 seconds.
These regular clicks were sometimes interrupted by
pauses of one to several minutes, or by a rapid
series of clicks that can sound like a creak. We
think that the regular clicks are used to acoustically
detect the sperm whales' prey. Do pauses and
creaks indicate that the whale has found potential
food?

After a bout of apparent foraging, which may
last from 5 to 30 hours, the group of sperm whales
will slow, and more and more animals will be
visible at the surface. Instead of being spread over
several kilometers of ocean, the whales begin to
cluster. We may see breaches (leaps from the
water) or lobtails (thrashes of the flukes onto the
water surface); we hear fewer of the rhythmic
"feeding" clicks, and more "codas." (Codas, such as
those analyzed by William A. Watkins of the
Woods Hole Oceanographic Institution, are
patterns of clicks apparently used for

Sperm whale calf with two adults photographed underwater. (Photo by Linda Weilgart)

communication between sperm whales.)

At the culmination of this clustering
behavior, the whole group of 20 or so sperm
whales may be gathered in a compact mass at the
surface. The whales lie quietly, sometimes a meter
or two apart. These "social" times can last
anywhere from 1 to 8 hours. As the clustering
ends, flukes are raised, and the whales
recommence their commute between the 410
meter depths for food and the surface for air.

The combined effect of about 15 sperm
whales at depth, each clicking once every 0.5 to 1
seconds sounds rather like radio static through a
hydrophone. But this blur of clicks was the beacon
that allowed us to follow a group of sperm whales
at night, and at other times when we could not see
them. Using a directional hydrophone, which can
detect sperm whales at about 10 kilometers, we
could tell the bearing and approximate range to the
whales, and thus follow them.

But sometimes, in the midst of the
cacophony of a feeding group of sperm whales, we
would hear a deeper, louder, and more ringing
click, repeating every 4 to 8 seconds rather than
0.5 to 1. These were apparently the "slow clicks" of
the large, mature male sperm whales. When heard
from very close range, the powerful slow clicks
sounded to us much like a slammed jailhouse door

might sound to a new inmate. These slow clicks
could be the large males' equivalent of the 0.5 to 1
second "feeding" click of the females; a signal to
receptive females; a signal to other males of the
size, strength, or reproductive status of the male;
or, they might serve combined functions.

Harems in Question

During our 2!/2 month study in the Galapagos
between February and April 1985, we heard slow
clicks with increasing frequency, and, during
daylight, would sometimes see the huge male
sperm whales themselves. They joined the groups
of females that we were following for an average of
about 6 hours at a time. While within the group,
they seemed to behave much like the smaller
whales that they were accompanying: they showed
their flukes and dived; they would sidle up beside
a female if nearby at the surface; and during "social
times" they would lie quietly at the surface among
the other whales. Sometimes we saw two, and
once three, large males with a group of females,
but there were no signs of aggression among them.

We were able to photographically identify
seven large males, probably representing the
majority of those off the Galapagos during our
study. Thus, we were able to trace which groups of
females, and which other males, a particular male

Sperm whales can be individually identified from photographs of their flukes. Two pictures of each of three sperm whales
are shown.

associated with. We found no signs of preference:
an individual male associated with a variety of
groups of females as well as other males. In
particular, there was no indication that males
formed harems out of the groups of females, or
formed consistent "coalitions" with other males.
The males seemed to move between groups. Thus,

male behavior may be somewhat less rigid than
previously believed. A similar pattern of males
moving independently between groups of females
has been found with African elephants.

Reduced Number of Males?

If our observations are representative of the form

Large male sperm whale with females. (Photo by V. Papastavrou)

of sperm whale social organization at other times
and in other oceans, this could have considerable
bearing on attempts to model the dynamics of
sperm whale populations. But another of our
observations could have even more significance:
the very small number of mature males present — a
maximum of 2 to 3 percent of the total Galapagos
population, rather than about 20 percent as several
assumptions about sperm whale natural history
would predict. This observation, and the possible
explanations for it, will need to be examined
through additional research. However, if there are
relatively fewer breeding males than pure
demography would predict, the relative reduction
of mature males, which have born the brunt of
recent whaling, may have an impact on the
probability of a female becoming pregnant. This
would lower the birth rate and lessen the ability of
the populations to recover from whaling.

For the moment this is speculation. We
need more information about the Galapagos sperm
whales. We need to know whether the females
that we observed form a discrete "Galapagos

stock," as some scientists believe, or whether
whales from a wider area of the Pacific use the
Galapagos from time to time. We need to know
the seasonality of breeding more accurately, and
whether our other observations of the behavior of
sperm whales are generally valid.

I Know Him Not

I am writing this during our second season (1986-
87) "on the Galapagos grounds." We plan to spend
a total of 7 months here, and then many more
analyzing the data back in our laboratories. By
then, we should know considerably more about
the Galapagos sperm whales. But, as to their "true
nature," I tend to agree with another visitor to the
Galapagos with an interest in sperm whales,
Herman Melville, who wrote of the central figure in
Moby Dick: "I know him not and never will."

Hal Whitehead is an Assistant Professor of Biology at
Dalhousie University, Halifax, Nova Scotia.

Huddled, or clustered "social"
sperm whales. (Photo by Tom
Arnbom)

Marine Iguanas:

Living on the Ocean Margin

by Andrew Laurie

I he marine iguanas of the Galapagos resemble
prehistoric reptiles. But, despite their formidable
dinosaur appearance, recent studies have
documented their general decline in numbers, and
sensitivity to environmental pressures. Of particular
interest are the responses of the populations to the
1982-83 El Nino event.

Origins and Characteristics

At the height of the Mesozoic, 200 million years ago,
there were many marine reptiles; now only a few
species of turtles, sea snakes, and crocodiles live in
the sea. A handful of species of terrestrial lizards
have secondarily become adapted to feeding, often

Land iguana (Conolophus subcristatusj feeding on Fernandina Island. (Photo by Jui De Roy, courtesy Galapagos National Park
Service)

opportunistically, on the ocean margin. The gecko,
Lepidodactylus woodfordi, hunts crabs just above the
high-tide zone in the Philippines, and iguanid lizards
of the genus Ctenosaura do the same in California
and Central America. On the Columbian island of
Malpelo, the lizard Anolis agass/z/, and the skink,
D/p/og/ossus hancocki, live partly or entirely on
crustaceans caught in the intertidal zone. Similar
behavior has been described by Hans Fricke for the
skink, Cryptoblephasus butoni, on the island of Nossi
Be near Madagascar, where they feed on insects,
crustaceans, and fish.

The marine iguanas of the Galapagos are the
descendants of iguanas, perhaps very similar to the
green iguana (/guana /guana) of the South American
mainland, which arrived in the islands less than 3
million years ago, probably on rafts of vegetation
swept out to sea from the rivers of the continent.
There are also two species (Brachylophus spp.) of
iguanas in Fiji, on the other side of the Pacific. Here
again, they are thought to have arrived there by

rafting from America where all the other 27 species
of iguanine lizards live.

However, the marine iguana, Amblyrhynchus
cristatus, is the only truly marine lizard in the world,
and has been equipped by natural selection within
the surprisingly short time of 2 to 3 million years with
flattened tails for swimming and long claws for
clinging to barnacle-covered rocks in the intertidal
surf. The marine iguanas are well-equipped for both
grazing at low tide on the abundant seaweed, and
for diving. While the iguanas can spend long periods
feeding underwater, the water in the Galapagos,
despite being on the equator, is often cold — mostly
because of the influence of nutrient-rich waters of
the Humboldt Current. Therefore, the iguanas must
warm their reptilian bodies by basking on the black
lava rocks of the shoreline.

The marine iguana is widely distributed
throughout the archipelago. The highest
concentrations are on the western islands. The
iguanas feed on soft-bodied, macrophytic marine

algae, either diving for them or grazing on exposed
intertidal rocks at low tide. The amount of feeding
beyond the tidal range varies between sites. Larger
individuals generally do more sub-tidal feeding,
while the smaller ones are restricted to the intertidal
zone.

Adult body size varies considerably between
islands, and (less so) between sites on the same
island: there is a ten-fold difference in adult male
body weight between the two extremes, Isabela and
Genovesa. Adult males range from 60 to 140
centimeters in length, and from 1.2 to 12.3 kilograms
in weight. The snout is short, and is the feature from
which the generic name (amblys = short, rhynchos =
snout) has been derived. The basic coloration is
black to dark grey, becoming light grey on the belly.
During the breeding season, and on some islands
throughout the year, the sides of the body and mid-
dorsal parts of the head and legs acquire a red and
greenish coloration.

Males defend mating territories for up to three
months during the annual breeding season. Females
lay 1 to 6 eggs in burrows dug 30 to 80 centimeters
deep in sand or volcanic ash, often up to 300 meters
or more inland. The eggs are left unattended and the
incubation period is approximately 95 days. There
have been big changes in the distribution and
abundance of marine iguanas during the last 1 50
years, and some of the early references to marine
iguanas in the literature enable interesting
comparisons to be made with the present situation.

Some History

The first reference to marine iguanas is by Fray
Tomas de Berlanga who found "many iguanas that
are like serpents," when he discovered the islands in
1 535. The buccaneers of the late 1 7th century wrote
in their journals of enormous numbers of iguanas,
which they often collected in large quantities for
food.

By the late 18th century the whalers were
frequenting Galapagos waters: Captain Colnett
writing in 1 798 regarded the "sea guana" as "the
most extraordinary animal" of Galapagos, and noted
that it "abounds in all these isles" and "goes to sea in
herds, a fishing"!

Porter, writing in 1822, also found "myriads of
guanas, of an enormous size and the most hideous
appearance imaginable." He "first supposed them
prepared to attack," and then tells how he and his
crew soon "discovered them to be the most timid of
animals, and had, in a few moments knocked down
hundreds of them with our clubs, some of which we
brought on board and found to be excellent eating."
This was on the south coast of Isabela in a place
where few iguanas remain today.

Marine iguanas were collected extensively
from many of the islands of the archipelago by the
various scientific expeditions of the late 18th and
early 19th centuries. Henry Blake noted an
abundance of marine iguanas in Tagus Cove, Isabela
in 1872; today hardly any remain there.

Reasons for the Decline in Numbers

Iguanas are patchily distributed throughout the
islands, with enormous concentrations in some areas

and very low densities in others. Population
compositions also vary greatly, with juveniles very
rare or totally absent in some areas. The iguanas
prefer the exposed, southern coastlines to the
sheltered, northern coastlines. The greatest
concentrations of marine iguanas occur where there
are shallow reefs and extensive intertidal zones.

Suitable nesting sites are also important in
determining the distribution of the major colonies,
but females may travel considerable distances to
nest.

Predation by introduced animals has a
great influence on abundance and population
composition. Introduced dogs, cats, rats, and pigs all
eat marine iguanas or their eggs. At Cabo Berkeley
and Muneco on northern Isabela, cats take almost
every hatchling before they reach one year of age,
so there is effectively no recruitment to the adult
population. At Caleta Webb on southern Isabela,
dogs were taking at least 27 percent of the
population each year until a poisoning campaign
eradicated them from the site in 1983.

Effects of the 1982-83 El Nino

The principal oceanographic event of the century in
the Galapagos, the 1982-83 El Nino/ENSO event,
had a profound effect on the iguanas. There was a
distinctly increased mortality rate, and those iguanas
that lived showed clear morphological, physiological,
and behavioral changes.

During 1983, an unusually high mortality of
marine iguanas was observed in populations on all
the islands of the archipelago. A major change in the
marine algal flora was observed during the same
period and abnormally high sea-surface
temperatures and sea-levels associated with the
El Nino/Southern Oscillation event were recorded
from November 1982 until July 1983. The abnormal
iguana mortality began in December 1982 and
continued until August 1983.

Every coastline was strewn with dead or dying
iguanas — thin, emaciated creatures half their former
weights — the survivors tried to eke an existence
from eating sea lion feces, crabs, or corpses of other
iguanas. The animals died of starvation, at least
partially attributable to their inability to digest an
invading species of alga, Griffordia mitchelliae, not
previously recorded in Galapagos. This invading
species was later again replaced by the normal food
species of red and green algae (for example,
Gelidium, Centroceras, Spermothamniutn, and Ulva
species).

In 1984, there was hardly any nesting, since
the survivors were in very poor condition. But, since
then the iguana population has shown a marked
recovery, with increased rates of growth, survival,
and reproduction.

In the course of our research, we monitored a
number of the principal effects of the event on the
life history of the iguanas:

Changes in annual rates of mortality. Annual
mortality rates before the 1982-83 El Nino varied
from 4 percent in adult females to 46 percent in
hatchlings. They shot up during the El Nino period to
53 percent in adult females, 63 percent in adult

100

3
If)

(fl

3
0)

50 H

1978-79
Hatchlmgs

'980Ha'ch/,ngs
I§8l_Hatchiings

i i i i i i

1981 1982 1983 1984 1985 1986
Year

Figure 1 . Cumulative percentage survival for various sex and
age classes of iguanas by year from 1 98 1 through 1 986.

males and 85 percent in hatchlings. These rates have
since returned to approximately pre-El Nino levels.
Figure 1 shows the cumulative percentage of
survivors in each year for different sex and age
classes. The 1985 hatchlings suffered high first year
mortality (about 60 percent) when compared with
the pre-El Nino first year mortality of the 1981
hatchlings (about 46 percent). But, as there were
approximately 1.8 times as many 1985 hatchlings as
the average annual production between 1981 and
1 983, there were more surviving yearlings of the
1985 cohort (year-class) than of the 1981 cohort.
This means that population density has remained
high for juveniles, but has fallen sharply for adults.

The high El Nino mortality is now being
compensated for by increased growth rates, earlier
breeding, more frequent breeding, and larger clutch
sizes. There was selection for large body size and
weight among juveniles in 1 983, and there were also
significant differences in their 1981 weights between
juveniles that subsequently survived and those that
did not survive the 1982-83 El Nino.

Growth rates. We have used the snout-vent
length (SVL) of the iguana as our main measure of
size and growth. Figure 2 shows the growth rates of
females as the mean annual increase in SVL plotted
against the SVL at the beginning of the year, for each
year since 1981. In other words, it shows the mean
growth rates for each size class of iguana. Growth
rates were clearly depressed during El Nino, to zero
in the larger animals, but rapidly increased
afterwards and are now gradually decreasing again
toward pre-El Nino levels. Immediately after El Nino,
one and two year-olds were growing at about twice
pre-El Nino rates.

Figure 3 shows the growth-curves for each
hatchling cohort from 1980 to 1986. The highest first

year growth was recorded in 1985 hatchlings; it has
decreased for the 1986 cohort. There was
considerable overlap in size distribution between the
1981 and 1982 cohorts and the 1982 and 1983
cohorts during their early growth.

Figure 4 shows the mean predicted growth in
SVL for both sexes, based on the 1981-82 and the
1985-86 data on annual increments in SVL for each
1 centimeter size class and both sexes. It shows a
clear shift in the curve to the left, so that both males
and females could be expected to reach the mean
size of 1981 breeding animals about two years earlier
than in 1981. Increased adult growth rates also
indicate a greater maximum attainable size.

Breeding age, nesting frequency, and clutch
size. A host of changes in reproductive rate followed
the period of stress and the associated recovery. As
predicted, one of the consequences of the increased
post El Nino growth rates was that females started
breeding at 2.5 years of age in 1985, compared with
4.5 years or more in previous years. Furthermore,
before El Nino, females nested approximately every
two years, with about 40 percent of females nesting
each year. Immediately after El Nino, there was
hardly any nesting on most islands, but since then
about 85 percent of females have nested each year,
and the mean clutch size has risen from 2 to 3 on
Santa Fe (Table 1), and from 3 to 6 on Isabela. In the
1985-86 nesting season, 1981 hatchlings nested for
the first time, but 1982 and 1983 hatchlings also
nested for the first time, both cohorts having attained
adult size by very fast growth after El Nino.

Changes in body condition. In a method
familiar to nutritionists and dieters, we prepared a
measure, or index, of body weight relative to body
dimension. This provides a measure of "fatness" or

1984-85

1985-86

1986-87

10 15 20 25 30

Snout-vent Length in April /cm

Figure 2. The average annual increase in snout-vent length
(SVL), based on the SVL at the beginning of the year (April) for
female iguanas, 1981 through 1987.

E
o
\

30-

25H

« 20

«= 15

1981

1 ' i '
82

' i ' i
83 84

Year

86 8

Figure 3. Growth curves (or each iguana year-class from 1 980
through 7986. Vertical error bars represent 1 standard
deviation.

40-,

Males

1 2 3 4 5 6 7 8 9 10 11 12 13 14
Age / Years

Figure 4. Average predicted growth in snout-vent length (SVL)
for both males and females, based on 1981-82 data and
1985-86 data. The average size of a 1981 nesting female was
24.5 centimeters SVL, a 1981 breeding male, 31.3
centimeters SVL. The smallest 1981 nesting female was 22.5
centimeters SVL, and the smallest 1981 breeding male was
28.0 centimeters SVL.

"thinness." In our case, we used weight divided by
SVL cubed: Weight/SVL3. This yielded a suitable
measure of the general condition of the iguanas
across age and sex classes.

Figure 5 shows the changes in mean
condition of adult iguanas (greater than 23
centimeters SVL) on Santa Fe over the 7 years 1981-
1987. Results for other islands are similar. The clear
trough in 1983 when many animals lost almost 50
percent of their body weight before either dying or
recovering, was followed by a sharp rise in condition

to well above the 1981 level, and then a return
towards that level.

Population Regulation

Over time, and through both major and minor
environmental perturbations, two factors seem to be
most important in determining the population size of
the iguanas: food and predation.

Food supply. The effects of the disappearance
of the preferred algal food species during the 1982-
83 El Nino were devastating, but such severe events
are infrequent, probably occurring not more than
once a century. There is evidence that the last event
of comparable magnitude to the 1982-83 El Nino
occurred in 1877-78. There are no reports of
widespread iguana mortality during previous, less
severe, El Nino events and although Ciffordia
mitchelliae was recorded again in early 1987, during
a minor El Nino event, and there was a reduction in
standing crop of red algae, no increased mortality
has yet been observed. However, condition and
growth rates are likely to be decreased by even
minor El Nino events, and thus lead to lower
reproductive output and possibly higher mortality, so
all El Nino events may be important in population
regulation.

Predation. Although iguanas appear to be
food limited, predation on land by introduced
species has probably been the main cause of the
gross changes in distribution and abundance of the
species during the past 150 years. Man also caught
the animals for food, but has never predated the
species at a significant level over large areas. Even so,
there are reports from San Cristobal that iguanas are
killed occasionally for dog food despite legal
protection. Cats are the main culprits among the

70 -i

50 -

c 40
O

1981

84
Year

Figure 5. The "condition index," a measure of body weight
relative to body dimension, for adult iguanas on Sante Fe
Island between April 1981 and February 1987. Vertical error
bars represent 1 standard deviation.

introduced predators — and Isabela, San Cristobal,
and Santa Cruz the islands most affected.

By poisoning, dogs have been successfully
eradicated from certain coastal areas, although
populations remain in the highlands. Complete
eradication has proved impossible so far, and in the
case of cats it is probably an unrealistic aim at this
stage. Local control around nesting areas of marine
iguanas and other species is probably the best plan.
Elsewhere, in New Zealand for example, the control
of feral cat populations has proved exceptionally
difficult and expensive. The task on the Galapagos,
especially on Isabela, is much more difficult because
of the treacherous lava shores, along which it is often
difficult, or even impossible, to walk, let alone
operate a cat control program. A pilot project is
needed in locations where cat predation is
particularly serious.

The Galapagos Marine Resources Reserve

Although the predation on land is the most
immediate problem for the marine iguanas on
islands with introduced predators, the long-term
security of the food supply is obviously vital, and the
establishment of the Galapagos Marine Resources
Reserve an essential step for the long-term
conservation of the species. Totally dependent on
the red and green algae of the intertidal and upper
subtidal zones, iguanas are particularly vulnerable to
marine pollution. As the species composition of the
algal flora is important there may be pollutants
which, although not fatal to all algae lead to a
decrease in the availability of iguana food species.
Certain species could become poisonous after
absorption of toxic chemicals.

Large oil spills would surely be fatal. There
already have been reports of minor oil slicks. One
came ashore on the south coast of Santa Fe in
January 1986 and killed the algae on a small part of
the intertidal zone. There is a constant danger of
more spills and major accidents. Nesting beaches
also could be affected, and hatchlings might be
particularly at risk, being restricted by their size, to
feeding at the very top of the intertidal zone.

Much of the pollution which may affect
Galapagos waters, particularly that originating
outside territorial waters, will not be made any easier
to control by the establishment of the marine

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reserve. However, the legal liability of ship owners
and captains for pollution within the reserve should
be used, in addition to conservation arguments, to
encourage correct maintenance and safety
procedures, and all that is possible to reduce the risk
of such accidents. Iguanas are particularly vulnerable
because of their low reproductive rate. Every
precaution must be taken to avoid pollution of any
sort, and this includes provision of equipment in
Galapagos for fighting oil spills, for example,
detergent sprayers, at such times as they are
considered preferable to the oil.

With the establishment of the marine reserve
should come a systematic monitoring program for
the marine environment, so that pollution,
overfishing of fish and lobsters, and unacceptable
human disturbance can be detected quickly and
controlled. For instance, regulations must be
enforced to limit the ever increasing pollution of the
coastlines of even the most remote islands with
plastic — bags, bottles, and other containers — which
threaten many coastal species, including iguanas. In
some cases it may be necessary to ban the import of
certain pollutants — for example plastic cola
bottles — to the islands, particularly when suitable
alternatives are readily available.

/Andrew Laurie is a Research Associate at the Max-Planck-
Institut fur Verhaltensphysiologie, Seewiesen, West Germany.
He is also affiliated with the Department of Zoology, Large
Animal Research Group, Cambridge University, England.

Acknowledgment

This is contribution No. 391 of the Charles Darwin
Foundation.

Selected References

Cooper, ). E., and W. A. Laurie. 1987. Investigation of deaths in

marine iguanas (Amblyrhynchus cristatus) on Galapagos. I. Comp.

Path. 97: 129-136.
Eibl-Eibesfeldt, I. 1984. The large iguanas of the Galapagos Islands.

In, Key Environments: Galapagos, ed. R. Perry, pp 157-173. New

York: Pergamon Press.
Kruuk, H., and H. Snell. 1981. Prey selection by feral dogs from a

population of marine iguanas (Amblyrhynchus cristatus). 1. Appl.

Ecol. 18: 197-204.

Laurie, A. 1983. Marine iguanas in Galapagos. Oryx 27: 18-25.
Laurie, A. 1983. An ill wind for iguanas. New. Scientist 100: 108.
Laurie, A. 1983. Marine iguanas suffer as El Nino breaks all records.

Noticias de Galapagos 38: 11.
Laurie, A. 1984. Marine iguanas: aftermath of El Nino. Noticias de

Galapagos 40: 9-11.

The Urvina Bay Uplift:

A Dry Trek Through
a Galapagos Coral Reef

by Mitchell W. Colgan, and David L Malmquist

Strange sights are commonplace in the Galapagos.
Still, it is surprising to find a place like Urvina Bay,
where trees grow out of coral heads and land
iguanas roam where fishes recently swam. The
circumstances that produced this unusual meeting of
land and water provide us with a natural outdoor
laboratory to explore topics in paleoecology,
sedimentology, and marine biology.

On a Wednesday morning in September,
1835, the H.M.S. Beagle sailed up the Bolivar
Channel toward an overnight anchorage at Bank's
Cove. On board was Charles Darwin. As the Beagle
passed within sight of the shallow waters of Urvina
Bay, Isabela Island, the young naturalist unknowingly
bypassed a rich coral community, where at least one
colony measured more than 5 meters in diameter.
Indeed, Darwin found neither coral communities nor
coral buildups during his 5 weeks in the Galapagos.
Unlike the pivotal role they played in his theory of
evolution, the islands did not play a part in Darwin's
ideas about the formation of coral reefs and atolls.

In 1954, however, the floor of Urvina Bay was
suddenly thrust more than 6 meters above sea level
(Figures 1, 2, and 3). Now, 152 years after the Beagle
visit, it is possible to casually take a dry walk through
a coral community and visit a part of the Galapagos
unknown to Darwin.

The Uplift

Urvina Bay lies between the Alcedo and Darwin
volcanoes on the west-central coast of Isabela Island.
Before the 1954 uplift, the coastline of Urvina Bay
lay in the shape of a "7," with the rocky coast on the
western flanks of the Alcedo volcano forming the
vertical stroke, and the mangrove and sandy-
beached shore adjacent to the Darwin volcano the
horizontal. Where these two shorelines met was a
shallow bay that harbored a rich coral and algal
community.

Volcan

Alcedo
Urvina Bay "^K (1128m)

•0:20'

-0°25'

91°20

91°15

Figure 1 . Urvina Bay on the west-central coast of Isabela
Island, showing the pre-uplift coastline and the 6-meter
depth contour.

This all changed early in 1954, when magma
beneath the Alcedo volcano suddenly rose, thrusting
the shoreline of Urvina Bay 1.2 kilometers seaward,
exposing several square kilometers of its marine
community.* Although the movement of this magma
was originally thought to be a precursor to a 1954
eruption of the Alcedo volcano, Keith Howard, a
scientist at the U.S. Geological Survey, has found no
evidence for such an eruption.

* Other uplifts are present in the Galapagos. Almost all are
smaller and more localized, and/or occurred along more
steep-sided coastlines. The Urvina Bay uplift is unique
because it elevated a shallow bay, thereby exposing an
extensive area.

Figure 2. Urvina Bay in 1946, eight years before the uplift.
The arrow points to a white, carbonate-sand beach, on either
side of which grew mangrove trees.

Figure 3. The same area in 1968, 15 years after a magmatic
disturbance uplifted Urvina Bay. Again, an arrow points to
the white, carbonate-sand beach, now more than a kilometer
from the water. (Photo by Tom Simkin)

The shallow bay seen by the Beagle's crew
had disappeared; in its place was a white, rocky
coastline. Jack Couffer, a member of a Walt Disney
crew filming in the Galapagos, was the first to report
the event (see "The Disappearance of Urvina Bay,"
Natural History, 1956). He found "in every niche and
crack and cave in the rock . . . the skeleton of some
sea animal — a crab, a starfish, a sea urchin, a fish. In
the depression below the rocks, the dried remains of

lobsters, sea turtles, and marine worms lay
mummified in the sun."

The very presence of these vertebrate and
invertebrate remains indicates that the uplift
occurred rapidly, perhaps overnight. Moreover, the
distribution of the remains suggests that most of the
water retreated from the uplift by percolating
downward through the porous aa lava (aa, from the
Hawaiian "to hurt" is fractured, jagged lava), rather
than by flowing overland. For instance, in
depressions seaward of the original mangroves are
the sun-bleached bones of more than 30 sea turtles.
In contrast to the remains of recently dead sea turtles
scattered by currents on the modern beach, these
skeletons occur in concentrated piles.* Unaltered,
large-scale sand ripples and the preservation of the
original angles of repose on Urvina Bay's former
beaches also give evidence of the rapid downward
flow of water. If the uplift had occurred over time,
we would expect to see a series of terraces on the
beaches, marking the level at which they temporarily
equilibrated to the new sea level.

The rapid uplift and drainage preserved the
integrity of the community and its sediments. Now,
some 33 years after the event, we can study subtidal
marine communities without getting wet (except
during the heavy rains of an El Nino year!).

The Lay of the Land

The dark basaltic lava of the Darwin Volcano is
Urvina Bay's foundation; its complex flow patterns
and topography of collapsed lava tubes left a
structural legacy that controlled the settlement and
growth of the bay's marine community. Variations in
water depth, degree of wave exposure, and
irregularity of the porous lava provided many
microhabitats where coral, calcareous algae, and
mollusks settled and grew. On the uplift, the remains
of these organisms produce a white-carbonate line
that delineates the old mean-low-water mark. This
cuts across the black basalt like a milky bathtub ring.

On the upraised shelf, there is a seaward
progression from landlocked sandy beaches and
mangrove swamps to areas underlain by deposits of
carbonate sand up to 2 meters thick. Isolated,
incipient coral reefs dot this area. More than a
kilometer from the stranded mangroves of the
uplifted shoreline, new mangroves grow and
flightless cormorants nest in an intertidal area only
33 years old. Along the new shore, beaches have
developed, and sea turtles now come to nest.

Coral Communities

Many atolls, fringing, and barrier reefs of the Pacific
arose from volcanic platforms similar to Urvina Bay.
But the origin of these reefs is now shrouded under
thick layers of carbonate. Urvina Bay's youthfulness

* As the reef section was uplifted, the turtles were trapped
in pools. Because the area was quickly elevated above sea
level, it was removed from the influence of waves and
currents, which would have reworked and scattered the
skeletons.

A tree grows from the center of a massive star coral, Pavona clavus. (Photo by Mitchell Colgan)

and its abundant exposures provide a rare
opportunity to document the initial steps in the
growth and expansion of a coral reef.

Physical and biological forces control the
composition and distribution of sediments shed by a
coral community during its growth. As the
community develops, the sediments change. Cores
reveal the expected — an overall upward increase of
carbonate deposition as the shelf matured.
However, layers with high concentrations of basaltic
sand interrupt the carbonate sequence. These
intervals of reduced reef growth and decreased
carbonate production may be linked to climatic
changes.

Red calcareous algae were the first colonists
on the barren basalt; these were followed by corals.
Among the earliest was the sand coral Psammocora
(Plesioseris) superficial is. In the central portion of the
uplift, where large colonies of the finger coral
Pocillopora damicornis now predominate, trenches
reveal an early shift in the community from the
smaller and slow-growing Psammocora to the larger
and faster-growing Pocillopora.

Although the eight hermatypic (reef-building)
coral species that inhabited Urvina Bay occur in
nearly every subenvironment on the uplift, they are
concentrated in only 10 locations. Three species—
the star coral Pavona clavus (Figure 4), the head coral
Porites lobata, and P. damicornis — have produced
five monospecific reefs. At first thought, these stands
might reflect distributional constraints brought about
through competition or genetics. But, with few
exceptions, competition between corals is not
apparent. Moreover, as defined by inferred water
depth and current exposure, most of the corals lived
in similar environments. Without any outstanding
environmental constraints on their distribution, the
segregation of corals on the uplift may simply reflect
chance settlement.

Because the Galapagos are bathed in
relatively cool waters, they are a marginal reef-
growing environment. Once coral larvae settle there,
the harsh conditions may limit their sexual
reproduction. Corals in the Galapagos compensate
by reproducing asexually, through fragmentation and
regrowth. For example, the counting of growth
bands on a giant colony of P. clavus by Gerard
Wellington, an Assistant Professor of Biology at the
University of Houston, shows this coral to be at least
350 years old. Nevertheless, except for fragments at
the coral's base, there are no other individuals of P.
clavus within 50 meters of this coral. Thus, for a third
of a millenium this coral grew and survived, but
apparently could not sexually reproduce. Because
asexual reproduction through fragmentation limits
the ability of corals to disperse, Galapagan corals
remain clustered in the random areas where
colonization originally occurred.

Fragmentation, and hence asexual
reproduction, of corals in the Galapagos is
accelerated by bioerosion. We are thus faced with a
contradiction worthy of Orwell's 1984: erosion is
growth. At Urvina Bay, bioeroders are common,
ranging from fishes to sponges, but the most
conspicuous eroder is the club-spined sea urchin

Eucidaris thouarsii. Coral colonies on the uplift are
often so heavily infested with Eucidaris borings that
they are nothing but thin shells arching over a hollow
interior. But like the saplings that spring from the
stump of a fallen tree, the fragments produced by
bioeroders commonly fall around the perimeter of
the parent colony, where they continue the colony's
growth. Many of Urvina Bay's corals apparently
reproduced most successfully through this type of
asexual fragmentation.

Galapagos reefs are smaller and less diverse
than most Pacific reefs. Nonetheless, understanding
their development should throw light on the origin
and history of larger oceanic reefs in the Pacific.

Climatology

In the Galapagos, as described by P. W. Glynn and
G. W. Wellington (see Selected Readings), climate
and water temperature play important roles in
determining the distribution of reefs and their rate of
development. Seasonal pulses of cold, upwelling
water, concentrated in the southwestern sector of
the archipelago, largely restrict reef growth to the
northeastern portion of the islands. Located in the
lee of Fernandina Island, Urvina Bay is somewhat
protected from these upwelling waters, and thus
corals can grow on the western coast of Isabela.

The 1982/83 El Nino-Southern Oscillation
(ENSO) event warmed the seas in the eastern Pacific
far above normal for many months, severely harming
Galapagos marine life and adding a new climatic
factor into the equation of the islands' reef
development (see article page 42). This El Nino killed
vast numbers of corals in the Galapagos and nearly
caused the localized extinction of P. damicornis.
According to Peter Glynn, a Professor of Biology at
the Rosenstiel School of Marine and Atmospheric
Science of the University of Miami, such severe El
Nino events may be important forces in structuring
the marine communities of the eastern Pacific. The
repeated thermal stresses suffered during El Nino
years may be partially responsible for the small size
of Urvina Bay's coral reefs.

The impact of El Nino events on eastern
Pacific marine communities depends on the
frequency with which the events occur in relation to
the lifetime of the communities. However, the long-
term record of the periodicity and ferocity of ENSO
events is poorly known. Our understanding of
marine and terrestrial community development
under El Nino stresses thus remains incomplete.
More importantly, a long-term El Nino record would
help planners prepare for future El Nino events.

Urvina Bay may provide us with this record of
El Nino events. Because El Ninos change the
temperature of the sea, they also alter the ratio of
stable-oxygen isotopes that corals incorporate in
their skeletons during growth. By analyzing the
isotopic signal recorded in the calcium carbonate
(CaCO3) skeleton of the giant P. clavus, the thermal
condition of the ocean at the time of the coral's
growth can be reconstructed. Working with Glynn
and others, we extracted a 4-meter core from this
coral. The core represents at least 350 years of time,
extending the record of El Ninos to roughly 1600.

This delicate duster of Pavona
elegans shows the exquisite
preservation of the marine
community on the uplift.
(Photo by Mitchell Colgan)

The isotopic analysis of the core is being done by
Wellington and Rob Dunbar, an Associate Professor
of Geology at Rice University.

Taphonomy and Paleoecology

Because the history of life on earth can only be
reconstructed through fossil evidence,
paleontologists must know what information is lost,
and what retained, as a once living community
passes into a fossil state. Urvina Bay is a natural
laboratory in which to study this passage. Here, we
can compare the dead community on the uplift to
the living community in the water, and examine the
changes that take place as a once living community
passes into an assemblage of fossils.

Because organisms with durable hardparts are
more likely to be preserved than those without, the

incomplete fossil record is biased. For example, even
though insects are by far the most abundant and
varied of living organisms (with some 800,000+
species), they are rare as fossils. The habitat of an
organism also helps dictate the likelihood of its
preservation. Organisms that live or die in
environments where deposition prevails are more
likely to be preserved than are those inhabiting
environments of erosion. The fossil record is thus
strongly skewed toward marine organisms with
durable skeletons.

It was not until 1940, when the Soviet
paleontologist J. A. Efremov initiated the science of
taphonomy (the study of fossilization), that
paleontologists began to fully understand how
important the imperfection of the fossil record is to
their science. Taphonomy, from the Greek root

'igure 4. An incipient reef of
3avona clavus fragments.
rhese fragments were shed by
in older colony that bioeroders
destroyed before the uplift
occurred. (Photo by Mitchell
lolgan)

"taphos," for burial, studies the passage of organisms
from the biosphere to the lithosphere. As such, it
bridges the gap between biology and paleontology.

Common sense dictates that a fossil
community retains less information than its living
predecessor: paleontologists commonly speak of
"taphonomic loss." At Urvina Bay, this holds true for
certain taxa. Though we searched the entire area of
the uplift in detail, we found the remains of only 4
fish, 1 marine iguana, and 5 sea stars. Curiously,
although they were reported in early descriptions of
the uplift, we found no lobsters. Unlike the early
paleontologists in the Arctic who reportedly ate
frozen mammoth while drinking cocktails chilled by
Pleistocene ice, we were unable to dine on
sunbaked "langosta."

Without allowing for the obvious taphonomic
loss that removed these taxa from the fossil record,
we would certainly reconstruct a community
different from the one visible to the snorkeler in the
present-day waters of Urvina Bay. Parrotfish,
wrasses, blennies, the ubiquitous damselfish, and the
endemic marine iguana are important shapers of the
structure of the modern community. Disregarding
taphonomic loss, they would be only minor
components of our reconstruction.

The preservation of most hard-shelled groups
at Urvina Bay, however, is exquisite. Individuals of E.
thouarsii are found on the uplift just as they occur in
Urvina Bay today. They occupy holes of their own
making in basaltic boulders and coral heads, with
their jaw structure, the Aristotle's lantern, still
articulated inside the urchin. The jaw commonly
projects from the mouth, where its five teeth rest
against the once algal encrusted but now barren
surface. The urchins' spines lie in perfect halos
around the skeletal test, further evidence that the
uplift was rapid and that subsequent current activity
was negligible.

Also beautifully preserved on the uplift, and
extremely abundant, are gastropods. These are
commonly found crowded in depressions. Evidently,
as seawater in small pools and fissures evaporated
after the uplift, the snails concentrated in these tepid
and saline pockets, finally dying.

Even such delicate forms as sponges survived
the uplift. These animals occur on the underside of
basaltic cobbles and boulders (a lifestyle described
by the term cryptic) — some retain their original form
and surface texture. Other common cryptic animals
found are chitons, corals, serpulid worms, and
bivalves.

For hard-shelled and cryptic organisms then, a
curious pattern exists at Urvina Bay. Contradicting
paleontology's golden rule that the present is the key
to the past, at Urvina Bay, the past of 33 years ago is
a better key to the present than is the modern
marine community. Instead of taphonomic loss,
there has been taphonomic gain.

There are three main reasons for this reversal.
First, try as we might with snorkels, wetsuits, masks,
and flippers, humans are terrestrial, not marine,
mammals. Unlike marine researchers, we can, on the
uplift, submerge ourselves in our work without
holding our breaths for the results. Visibility on the
uplift is fantastic; we sometimes had "five volcano

days" when Alcedo, Darwin, Cerro Azul, Sierra
Negra, and Fernandina volcanoes were visible from
our study site — the farthest is 60 kilometers away.
Moreover, there is no surge or swell on the uplift
and we, rather than the Galapagos shark, are at the
top of the food chain.

Second, the carpet of fleshy algae that hides
so much life in the water is absent on the uplift,
revealing large numbers of gastropods, bivalves, and
arthropods that would otherwise go undetected.

Third, nocturnal, infaunal, and cryptic animals,
which are visible only with great effort to the marine
biologist, are on the uplift clearly exposed to the
equatorial sun.

Because of the exquisite preservation of the
hard-shelled taxa at Urvina Bay, our future research
there will concentrate on answering the question:
"How much information about an original living
community can a paleontologist garner based on a
complete representation of the hard-shelled taxa of
that community?" That is, even with no loss of
skeletal information, what can a paleontologist say
about the entire living community based on its
skeletal record alone? At Urvina Bay, the
juxtaposition of comparable fossil and living
communities puts the answer to this question within
our grasp.

Mitchell W. Colgan and David L Malmquist are doctoral
candidates in Earth Sciences at the University of California,
Santa Cruz. Both are conducting thesis research at Urvina
Bay, Galapagos, Ecuador, under the supervision of Leo F.
Laporte.

Acknowledgments

This research was funded by NSF grant EAR-8508966. The
Ecuadorian Ministero de Agriculture y Cranaderia and the
Departamento de Parques Nacionales y Vida Silvestre, and
the Charles Darwin Foundation granted us permission to
carry out this study. Assistance in the Galapagos was
provided by Cunther Reck, Sylvia Harcourt, Henk
Kasteleijin, and the rest of the staff at the Charles Darwin
Research Station. Miguel Cifuentes and Ing. Humberto
Ochoa Cordova at Parque Nacional Galapagos provided
help without which this research would not have been
possible. We thank those who helped us at Urvina Bay:
Linda Anderson, William Anderson, Sain Chai Colgan, Rene
Espinosa, David Hollander, Margaret Liniecki Laporte,
Christa Sadler, and Tom Smalley. We also give thanks to the
following for their assistance and support: Juan Black of the
Charles Darwin Foundation, Cynthia Colgen, Jessica
Colgan, Sarah Gray, Gene Gonzales, Sarah Griscom, Keith
Howard, Melissa Malmquist, R. Larry Phillips, and Jerry
Wellington. Special thanks are due to Robert Garrison,
Peter Glynn, Leo F. Laporte, Clif Jordan, and Gene Shinn,
all of whom helped in the early formulation of this research.
Charles Darwin Foundation contribution No. 414.

Selected Readings

Couffer, J. C. 1956. The disappearance of Urvina Bay. Natural

History 65: 378-383.
Darwin, C. 1842. The Structure and distribution of coral reefs.

London: Smith, Elder & Co.
Clynn, P. W. 1983. Extensive "bleaching" and death of coral on the

Pacific coast of Panama. Environmental Conservation 10: 149-

154.
Glynn, P. W., and C. W. Wellington. 1983. Corals and coral reefs of

the Galapagos Islands. Berkeley: University of California Press.

Galapagos Seabirds

liven though Darwin's finches may be the
group of birds most symbolic of Galapagos
ecology, the nesting seabirds of the islands form
a unique community of their own. Three ocean
currents — the cold Humboldt, the warm North
Equatorial, and the cold, upwelling Cromwell-
converge at the Galapagos, and each has
brought a characteristic set of seabirds to the
islands. According to ornithologist M. A. Harris,
the archipelago now provides nesting sites for
14 migratory and five endemic species,
representing a total of about 750,000 seabirds.

The Humboldt Current was probably the
highway taken by the most unlikely bird to nest
in the equator-straddling. islands, the Galapagos
penguin, Spheniscus mendiculus. These
penguins have long since lost touch with their
nearest relatives on the Patagonian coast, and
so are endemic to the Galapagos. More than
10,000 of them live in many colonies on the
islands, banding together in large groups to fish
the colder waters sweeping the islands.

Keeping cool under the equatorial sun is
more than a matter of comfort for these
antarctic outcasts. Ornithologist D. Boersma
reported a case in which one penguin, while
shading its nest from the sun, became so
overheated that it took a life-saving plunge into
the sea. Unfortunately, by the time it returned
to its nest, the eggs had overheated and never
hatched.

The penguins have a non-flying partner
among the Galapagos seabirds in the flightless
cormorant, Nannopterum harrisi. These two
species are the rarest seabirds in the world. The
cormorant is the only one of the 29 cormorant
species unable to fly. It has even lost the
enlarged keel on its breastbone for the

Blue-footed boobies. (Photo by Tui De Roy, courtesy
National Park Service)

The Galapagos penguin. (Photo by Tui De Roy, courtesy
National Park Service)

attachment of flight muscles; yet after diving it
still spreads out its vestigial wings to dry, in
characteristic cormorant fashion. Although it
feeds closer to shore than any other Galapagos
seabird, Darwin failed to notice it during his
1835 voyage.

The other large endemic seabird is the
waved albatross, Diomedea irrorata. Weighing-
in at 3 to 4 kilograms, and with a wingspan of
up to 2 meters, it is the largest species of
albatross. It also is the only tropical species
among the 13 of the family.

Nesting pairs of Diomedea mate for life,
which can last up to 50 years. Their courtship
behavior is curious in that its most elaborate
display is at the end of the breeding season.
This involves a clattering, bill-circling dance.
During the dance, the bills slide easily over one

another, lubricated by an oily secretion. Chicks
hatch out of eggs weighing about 285 grams,
and are fed a predigested, oily mixture of squid
and fish by their far-ranging parents. The chicks
can take in as much as 2 liters of this at one
feeding, stuffing them to the point of
immobility.

The brown pelican, Pelecanus
occidentalis, while being one of the largest
Galapagos seabirds, is one of the smallest
species of pelican. Many visitors are awed by its
vertical, 50-foot dives into the sea. On these
spectacular sorties it collects up to 3 gallons of
water in its expandable bill, and filters out meals
of small fish. Croups of brown pelicans are also
impressive as they fly almost in unison, just
above the surface of the warm waters of the
North Equatorial Current.

^ -,-. • -

The brown pelican. (Photo by DJ.H. Phillips)

Those waters a/so bring the three species
of boobies to the Galapagos. They are said to
have been so-named by seafarers of long ago,
because of their strange appearance and
behavior. "Bobo" is Spanish for clown — and the
term "booby-hatch" comes from the birds' habit
of diving headlong off the bows of ships, in
pursuit of flying fish.

Most remarkable in appearance is the
blue-footed booby, Sula nebouxii. Because it
feeds close to shore, it is seen more often than
its relatives — the masked and the red-footed

boobies, each of which outnumber the blue-
footed. Although S. nebouxii does not build a
nest for its eggs, it still engages in a vestigial
nest-building routine. The bare ground where
the eggs are laid often gets dangerously hot. So
the nesting booby shades them with its body,
and keeps them on top of its foot-webs.

Another warm-water Galapagos species
is the dark-rumped petrel, Pterodroma
phaeopygia. It is the most endangered of the
nesting seabirds. In Hawaii, it is almost extinct
as the result of predation by feral cats, rats, and
dogs. In the Galapagos, these same animals
have pushed this petrel into a "very precarious"
situation, according to seabird ecologist
Malcolm Coulter.

The white-vented storm-petrel,
Oceanites gracilis, is a species typical of the
cold Humboldt Current. While Coulter
estimates that many thousands of them nest at
the Galapagos, to date no nesting sites have
been discovered.

Even though increasing numbers of
tourists visit the Galapagos each year, the
nesting seabirds seem little affected by their
influence. Visitors must be accompanied at all
times by a naturalist guide. The seabirds were
taken into account when the boundary of the
Galapagos Marine Reserve was drawn (see map
page 2). The Reserve extends 15 miles beyond
the outermost islands, insuring safe fishing areas
for boobies, petrels, and their neighbors.

-TMH

Odd-Ball Uses for ROVs

There's a new manual for entrepreneurs called
"Sunken Golf Ball Recovery." It's no problem to
make $700 a day retrieving golf balls from water
hazards, reports SubNotes in its April issue.

One man in Florida reportedly dons SCUBA
gear and hauls up 300,000 golf balls a year. At $7
a dozen, that's a cool $175,000 before taxes and
overhead. A southern California operator
supposedly can collect 78,000 balls in a month.
He sells them at the bargain-basement price of
just 25C apiece, so his monthly income from the
venture is $19,500. Not bad.

But, these entrepreneurs are missing the high-
tech angle to this enterprise. Some ocean
engineer should come up with a golf-ball
recovery tool and basket attachment for
remotely-operated vehicles (ROVs). This would
open the golf-ball recovery game to non-divers.
One would avoid the inconvenience of getting
wet, and pocket a $1,000 a day or so to boot. It
is time to stop foolin' with shipwrecks in
inhospitable places, and turn the technology
attention to the more mellow environs of the
local golf course. Isn't this what the American
dream is all about?

A Search for Unique Drugs

in the Galapagos
Underwater Environment

by Shirley A. Pomponi, and Susan van Hoek

Scientists from the Harbor Branch
Oceanographic Institution/SeaPharm Project
spent seven weeks this past winter cruising the
Galapagos Islands searching for new drugs from
marine organisms. Results from bioassays taken
aboard the institution's research vessel 5eward
Johnson indicated that possibly as many as 15
percent of the organisms collected have
anticancer, antiviral, antimicrobial, or
immunomodulatory activity. More extensive
testing is now under way in the Harbor Branch/

SeaPharm Project laboratories in Fort Pierce,
Florida.

Nearly 1,200 organisms were collected by
wading, snorkeling, SCUBA diving, and diving
inside the four-man, Johnson Sea-Link I
submersible. Of these samples, nearly 700 were
collected at depths from 350 feet to 2,630 feet
(near maximum diving range for the

Above, (he Johnson Sea-Link I. (Photo courtesy Harbor
Branch Oceanographic Institution)

Shirley Pompon/ and soft coral on a vertical wall at 70 feet. (Photo by John Reed)

submersible). The most abundant animals
collected were sponges, echinoderms (primarily
starfishes, sea urchins, and sea cucumbers), and
soft corals, though representatives of most
marine invertebrate phyla, as well as algae and
ascidians (sea squirts), were also collected.

The search for cures for such diseases as
cancer from marine organisms follows extensive
evaluations of land organisms. To date, only
about 20 clinically useful drugs are available for
treatment against cancer, and these are toxic
and have a limited range of activity. Scientists
are now searching underwater on the
presumption that the oceans may provide a vast
reservoir of untapped resources.

The Galapagos Islands were targeted as a
possible source for new drugs because the area
has a unique, relatively unexplored marine
environment. Some of the most interesting
collections were made west of Isabela and
Fernandina Islands, the youngest and most
active islands of the volcanic chain and the site
of nutrient-rich, upwelled water.

Galapagos Flora and Fauna

Galapagos shallow water flora and fauna share
many similarities with that of the tropical eastern
Pacific, including the Gulf of California, although
a few species collected, particularly

echinoderms, are more closely related to
species in the Indian Ocean and western Pacific.
The intertidal areas are often described as
barren due to the large number of grazers and
predators, but cruise participant Richard C.
Brusca, Curator of Invertebrates, Los Angeles
County Natural History Museum, found the
rocky shores to be surprisingly rich in
invertebrates.

Relatively little is known about the Galapagos
deep sea fauna. The Johnson Sea-Link I
submersible, equipped with underwater video,
35-mm cameras, a manipulator arm, and a
suction device gave scientists new insights into
the deepwater communities that have been
traditionally sampled with trawling and dredging
rigs. Several new species of ascidians, or sea
squirts, were discovered. Cruise participant
Francoise Monniot, Curator of Marine
Invertebrates at the Paris Museum of National
History, reported that the sea squirts are
unusual for both their large size and their
taxonomic distribution. She said, "this may
change the general opinion about the diversity,
the origin, and the evolution" of deep water
ascidians.

Preliminary examination of more than 300
Galapagos sponges indicates several new
species were discovered. In addition, the deep

,*#•

A sea fan on a rocky volcanic slope. (Photo by lohn Reed)

A cluster of green sea urchins endemic to the Galapagos,
collected off Fernandina Island. (Photo by lohn Reed)

water "glass" sponges, or hexactinellids, were
much more diverse than those previously
studied at the same depths in the West Indies.
Collections of choristids — a group of sponges
with abundant, prickly, glass-like spicules—
show similarities with species collected in the
Caribbean. This may lead to new hypotheses
about the origin, evolution, and relationships of
that group of animals.

How the Samples Were Handled

After preserving a sample of each organism in
alcohol or formalin, a small piece was made into
an extract by grinding in a solvent mixture
containing alcohol. The rest of the organism was
preserved by freezing at minus 20 degrees
Celsius. Living organisms, as well as alcohol- and
formalin-preserved samples, were studied by
taxonomic specialists participating in the
expedition.

Extracts were tested by a team that included a
virologist, a microbiologist, an immunologist,
and a tissue-culture specialist. Of particular
interest is the ability of the extracts to inhibit
growth of cancer cells, viruses, bacteria, fungi,
and yeast, as well as their potential for
stimulating or suppressing the immune system.
Obvious applications of this research are the
development of drugs to treat cancers, viral
diseases, and diseases of the immune system,
such as AIDS.

The expedition was funded in part by a
contract between SeaPharm and the National
Cancer Institute, which will test the extracts
from deep water organisms against a number of
human cancers. This recently expanded interest
in marine organisms as a source of anticancer
drugs is based on discoveries of unique
biochemicals with pharmacological activity
produced by some marine plants and animals.

"Retro-spinoff," a Key By-Product

Although the primary goal of this research
mission was the discovery of unique chemical
compounds with pharmacological activity,
important marine biological discoveries were
also realized. Kenneth L. Rinehart, Natural
Products Chemist at the University of Illinois,
Director of Research at SeaPharm, Inc., and
Chief Scientist of the expedition, dubbed such
discoveries "retro-spinoff" or the gathering of
new information that will become a part of the
current pool of basic scientific knowledge.

Enough material was brought back from the
expedition to keep scientists busy for years,
especially with taxonomy (organism
identification) and systematics (life histories,
distribution, abundance, ecological,
phylogenetic, and evolutionary relationships
among organisms). Many of the samples
preserved as a reference collection will be
deposited at the Smithsonian Institution,
Washington, D.C., and at Harbor Branch.

SeaPharm, Inc., a pharmaceutical company
with headquarters in Princeton, New Jersey,
specializes in the discovery and development of
drugs from marine organisms. Harbor Branch
Oceanographic Institution, Inc., located on the
west bank of the Intracoastal Waterway
between Fort Pierce and Vero Beach, Florida, is
a not-for-profit organization dedicated to
research in the marine sciences and to the
development of tools and systems for
oceanographic research.

Shirley A. Pompon/ is a marine biologist specializing in
sponges, and a Senior Scientist with SeaPharm, Inc.
Susan van Hoek is Public Affairs Officer and a writer/
editor at Harbor Branch Oceanographic Institution, Inc.

The Voyage of the Beagle

EDITOR'S NOTE: What follows is an edited version
of Chapter 1 7 of The Voyage of the Beagle by
Charles Darwin. The celebrated naturalist landed
in the Galapagos on 16 September 1835 to begin
five weeks of collecting and observing, the results
chronicled in Chapter 17. In all, he visited four of
the major island in the group during his stay. As
this issue of Oceanus focuses on the new
Galapagos marine reserve, most of the material we
have deleted concerns Darwin's observations of
terrestrial fauna and flora.

CHAPTER XVII

Galapagos Archipelago
by Charles Darwin

SEPTEMBER 75th.— This archipelago consists of 10
principal islands, of which five exceed the others in
size. They are situated under the Equator, and
between 500 and 600 miles westward of the coast
of America. They are all formed of volcanic rocks; a
few fragments of granite curiously glazed and
altered by the heat, can hardly be considered as an
exception. Some of the craters, surmounting the
larger islands, are of immense size, and they rise to
a height of between 3 and 4,000 feet. Their flanks
are studded by innumerable smaller orifices. I
scarcely hesitate to affirm, that there must be in the
whole archipelago at least 2,000 craters. These
consist either of lava or scoriae, or of finely-
stratified, sandstone-like tuff. Most of the latter are
beautifully symmetrical; they owe their origin to
eruptions of volcanic mud without any lava: it is a
remarkable circumstance that every one of the 28
tuff-craters which were examined, had their
southern sides either much lower than the other
sides, or quite broken down and removed. As all
these craters apparently have been formed when
standing in the sea, and as the waves from the
trade wind and the swell from the open Pacific
here unite their forces on the southern coasts of all
the islands, this singular uniformity in the broken
state of the craters, composed of the soft and
yielding tuff, is easily explained.

current. Excepting during one short season, very
little rain falls, and even then it is irregular; but the
clouds generally hang low. Hence, whilst the lower
parts of the islands are very sterile, the upper parts,
at a height of a thousand feet and upwards, possess
a damp climate and a tolerably luxuriant
vegetation. This is especially the case on the
windward sides of the islands, which first receive
and condense the moisture from the atmosphere.

In the morning (17th) we landed on
Chatham Island, which, like the others, rises with a
tame and rounded outline, broken here and there
by scattered hillocks, the remains of former craters.
Nothing could be less inviting than the first
appearance. A broken field of black basaltic lava,
thrown into the most rugged waves, and crossed
by great fissures, is everywhere covered by
stunted, sunburnt brushwood, which shows little
signs of life. The dry and parched surface, being
heated by the noon-day sun, gave to the air a close
and sultry feeling, like that from a stove: we fancied
even that the bushes smelt unpleasantly. Although
I diligently tried to collect as many plants as
possible, I succeeded in getting very few; and such
wretched-looking little weeds would have better
become an arctic than an equatorial Flora. . . .

The Beagle sailed round Chatham Island,
and anchored in several bays. One night I slept on
shore on a part of the island, where black
truncated cones were extraordinarily numerous:
from one small eminence I counted 60 of them, all
surmounted by craters more or less perfect. The
greater number consisted merely of a ring of red
scoriae or slags, cemented together: and their
height above the plain of lava was not more than
from 50 to 100 feet; none had been very lately
active. The entire surface of this part of the island
seems to have been permeated, like a sieve, by the
subterranean vapours: here and there the lava,
whilst soft, has been blown into great bubbles; and
in other parts, the tops of caverns similarly formed
have fallen in, leaving circular pits with steep sides.
From the regular form of the many craters, they
gave to the country an artificial appearance, which
vividly reminded me of those parts of Staffordshire,
where the great iron-foundries are most numerous.
The day was glowing hot, and the scrambling over
the rough surface and through the intricate
thickets, was very fatiguing; but I was well repaid
by the strange Cyclopean scene. As I was walking
along I met two large tortoises, each of which must
have weighed at least 200 pounds: one was eating
a piece of cactus, and as I approached, it stared at
me and slowly walked away; the other gave a deep
hiss, and drew in its head. These huge reptiles,
surrounded by the black lava, the leafless shrubs,
and large cacti, seemed to my fancy like some
antediluvian animals. The few full-coloured birds
cared no more for me than they did for the great
tortoises.

23rd. — The Beagle proceeded to Charles
Island. ... In the woods there are many wild pigs
and goats; but the staple article of animal food is
supplied by the tortoises. Their numbers have of
course been greatly reduced in this island, but the
people yet count on two days' hunting giving them
food for the rest of the week. It is said that formerly
single vessels have taken away as many as 700, and
that the ship's company of a frigate some years
since brought down in one day 200 tortoises to the
beach.

slope, and, choked with dust, eagerly tasted the
water — but, to my sorrow, I found it salt as brine.

The rocks on the coast abounded with great
black lizards, between three and four feet long; and
on the hills, an ugly yellowish-brown species was
equally common. We saw many of this latter kind,
some clumsily running out of the way, and others
shuffling into their burrows. I shall presently
describe in more detail the habits of both these
reptiles. The whole of this northern part of
Albemarle Island is miserably sterile. . . .

flffS

153?

JO/ at ta^f

Cutaway view of H. M. S. Beagle. During the voyage, Darwin shared quarters with Captain FitzRoy. (Diagram courtesy of
Frank Sulloway)

September 29th. — We doubled the south-
west extremity of Albemarle Island, and the next
day were nearly becalmed between it and
Narborough Island. Both are covered with
immense deluges of black naked lava, which have
flowed either over the rims of the great caldrons,
like pitch over the rim of a pot in which it has been
boiled, or have burst forth from smaller orifices on
the flanks; in their descent they have spread over
miles of the sea-coast. On both of these islands,
eruptions are known to have taken place; and in
Albemarle, we saw a small jet of smoke curling
from the summit of one of the great craters. In the
evening we anchored in Bank's Cove, in Albemarle
Island. The next morning I went out walking. To
the south of the broken tuff-crater, in which the
Beagle was anchored, there was another beautifully
symmetrical one of an elliptic form; its longer axis
was a little less than a mile, and its depth about
500 feet. At its bottom there was a shallow lake, in
the middle of which a tiny crater formed an islet.
The day was overpoweringly hot, and the lake
looked clear and blue: I hurried down the cindery

October 8th — One day we accompanied a
party of the Spaniards in their whaleboat to a
salina, or lake from which salt is procured. After
landing, we had a very rough walk over a rugged
field of recent lava, which has almost surrounded a
tuff-crater, at the bottom of which the salt-lake lies.
The water is only three or four inches deep, and
rests on a layer of beautifully crystallized, white
salt. The lake is quite circular, and is fringed with a
border of bright green succulent plants; the almost
precipitous walls of the crater are clothed with
wood, so that the scene was altogether both
picturesque and curious. A few years since, the
sailors belonging to a sealing-vessel murdered their
captain in this quiet spot; and we saw his skull lying
among the bushes.

During the greater part of our stay of a
week, the sky was cloudless, and if the trade-wind
failed for an hour, the heat became very
oppressive. On two days, the thermometer within
the tent stood for some hours at 93 degrees; but in
the open air, in the wind and sun, at only 85
degrees. The sand was extremely hot; the

thermometer placed in some of a brown colour
immediately rose to 137 degrees and how much
above that it would have risen, I do not know, for it
was not graduated any higher. The black sand felt
much hotter, so that even in thick boots it was
quite disagreeable to walk over it.

The natural history of these islands is
eminently curious, and well deserves attention.
Most of the organic productions are aboriginal
creations, found nowhere else; there is even a
difference between the inhabitants of the different
islands; yet all show a marked relationship with
those of America, though separated from that
continent by an open space of ocean, between 500
and 600 miles in width. The archipelago is a little
world within itself, or rather a satellite attached to
America, whence it has derived a few stray
colonists, and has received the general character of
its indigenous productions. Considering the small
size of the islands, we feel the more astonished at
the number of their aboriginal beings, and at their
confined range. Seeing every height crowned with
its crater, and the boundaries of most of the lava-
streams still distinct, we are led to believe that
within a period geologically recent the unbroken
ocean was here spread out. Hence, both in space
and time, we seem to be brought somewhat near
to that great fact — that mystery of mysteries — the
first appearance of new beings on this earth. . . .

Of land-birds I obtained 26 kinds, all
peculiar to the group and found nowhere else,
with the exception of one lark-like finch from
North America (Dolichonyx oryzivorus). The other
25 birds consist, firstly, of a hawk, curiously
intermediate in structure between a buzzard and
the American group of carrion-feeding Polybori;
and with these latter birds it agrees most closely in
every habit and even tone of voice. Secondly,
there are two owls, representing the short-eared
and white barn-owls of Europe. Thirdly, a wren,
three tyrant-flycatchers (two of them species of
Pyrocephalus, one or both of which would be
ranked by some ornithologists as only varieties), a
dove — all analogous to, but distinct from, American
species. Fourthly, a swallow, which though
differing from the Progne purpurea of both
Americas, only in being rather duller colored,
smaller, and slenderer, is considered by Mr. Gould
as specifically distinct. Fifthly, there are three
species of mocking thrush — a form highly
characteristic of America.

The remaining land-birds form a most
singular group of finches, related to each other in
the structure of their beaks, short tails, form of
body and plumage: there are 13 species, which Mr.
Gould has divided into four sub-groups. All these
species are peculiar to this archipelago; and so is
the whole group, with the exception of one species
of the sub-group Cactornis, lately brought from
Bow Island, in the Low Archipelago. . . . The males
of all, or certainly of the greater number, are jet
black; and the females (with perhaps one or two
exceptions) are brown.

The most curious fact is the perfect
gradation in the size of the beaks in the different

species of Geospiza, from one as large as that of a
hawfinch to that of a chaffinch, and (if Mr. Gould is
right in including his sub-group, Certhidea, in the
main group) even to that of a warbler. . . .

Of waders and waterbirds I was able to get
only 1 1 kinds, and of these only three (including a
rail confined to the damp summits of the islands)
are new species. Considering the wandering habits
of the gulls, I was surprised to find that the species
inhabiting these islands is peculiar, but allied to
one from the southern parts of South America. The
far greater peculiarity of the landbirds, namely, 25
out of 26, being new species, or at least new races,
compared with the waders and web-footed birds,
is in accordance with the greater range which these
latter orders have in all parts of the world. We shall
hereafter see this law of aquatic forms, whether
marine or fresh water, being less peculiar at any
given point of the Earth's surface than the
terrestrial forms of the same classes, strikingly
illustrated in the shells, and in a lesser degree in
the insects of the archipelago. . . .

We will now turn to the order of reptiles,
which gives the most striking character to the
zoology of these islands. The species are not
numerous, but the numbers of individuals of each
species are extraordinarily great. There is one small
lizard belonging to a South American genus, and
two species (and probably more) of the
Amblyrhynchus — a genus confined to the
Galapagos Islands. There is one snake which is
numerous; it is identical. Of sea-turtle I believe
there are more than one species; and of tortoises
there are, as we shall presently show, two or three
species or races. Of toads and frogs there are none:
I was surprised at this, considering how well suited
for them the temperate and damp woods appeared
to be. It recalled to my mind the remark made by
Bory St. Vincent, namely, that none of this family
are found on any of the volcanic islands in the
great oceans. As far as I can ascertain from various
works, this seems to hold good throughout the
Pacific, and even in the large islands of the
Sandwich archipelago. . . . The absence of the frog
family in the oceanic islands is the more
remarkable, when contrasted with the case of
lizards, which swarm on most of the smallest
islands. May this difference not be caused, by the
greater facility with which the eggs of lizards,
protected by calcareous shells, might be
transported through salt-water, than could the
slimy spawn of frogs?

I will first describe the habits of the tortoise
(Testudo nigra, formerly called Indica), which has
been so frequently alluded to. These animals are
found, I believe, on all the islands of the
archipelago; certainly on the greater number. They
frequent in preference the high damp parts, but
they likewise live in the lower and arid districts. I
have already shown, from the numbers which have
been caught in a single day, how very numerous
they must be. Some grow to an immense size: Mr.
Lawson, an Englishman, and vice-governor of the
colony, told us that he had seen several so large,
that it required six or eight men to lift them from

the ground; and that some had afforded as much
as 200 pounds of meat. The old males are the
largest, the females rarely growing to so great a
size: the male can readily be distinguished from the
female by the greater length of its tail.

The tortoises which live on those islands
where there is no water, or in the lower and arid
parts of the others, feed chiefly on the succulent
cactus. Those which frequent the higher and damp
regions, eat the leaves of various trees, a kind of
berry (called guayavita) which is acid and austere,
and likewise a pale green filamentous lichen
(Usnera plicata), that hangs from the boughs of the
trees.

The tortoise is very fond of water, drinking
large quantities, and wallowing in the mud. The
larger islands alone possess springs, and these are
always situated towards the central parts, and at a
considerable height. The tortoises, therefore, which
frequent the lower districts, when thirsty, are
obliged to travel from a long distance. Hence broad
and well-beaten paths branch off in every direction
from the wells down to the seacoast; and the
Spaniards by following them up, first discovered
the watering-places.

When I landed at Chatham Island, I could
not imagine what animal travelled so methodically
along well-chosen tracks. Near the springs it was a
curious spectacle to behold many of these huge
creatures, one set eagerly travelling onwards with
outstretched necks, and another set returning, after
having drunk their fill. When the tortoise arrives at
the spring, quite regardless of any spectator, he
buries his head in the water above his eyes, and
greedily swallows great mouthfuls, at the rate of
about 10 in a minute. The inhabitants say each
animal stays three or four days in the
neighbourhood of the water, and then returns to
the lower country; but they differed respecting the
frequency of these visits. The animal probably
regulates them according to the nature of the food
on which it has lived. It is, however, certain, that
tortoises can subsist even on these islands where
there is no other water than what falls during a few
rainy days in the year.

I believe it is well ascertained, that the
bladder of the frog acts as a reservoir for the
moisture necessary to its existence: such seems to
be the case with the tortoise. For some time after a
visit to the springs, their urinary bladders are
distended with fluid, which is said gradually to
decrease in volume, and to become less pure. The
inhabitants, when walking in the lower district, and
overcome with thirst, often take advantage of this
circumstance, and drink the contents of the
bladder if full: in one I saw killed, the fluid was
quite limpid, and had only a very slightly bitter
taste. The inhabitants, however, always first drink
the water in the pericardium, which is described as
being best.

The tortoises, when purposely moving
towards any point, travel by night and day, and
arrive at their journey's end much sooner than
would be expected. The inhabitants, from
observing marked individuals, consider that they

travel a distance of about eight miles in two or
three days. One large tortoise, which I watched,
walked at the rate of 60 yards in 10 minutes, that is
360 yards in the hour, or four miles a day,—
allowing a little time for it to eat on the road.

During the breeding season, when the male
and female are together, the male utters a hoarse
roar or bellowing, which, it is said, can be heard at
the distance of more than a hundred yards. The
female never uses her voice, and the male only at
these times; so that when the people hear this
noise, they know that the two are together. They
were at this time (October) laying their eggs. The
female, where the soil is sandy, deposits them
together, and covers them up with sand; but where
the ground is rocky she drops them
indiscriminately in any hole: Mr. Bynoe found
seven placed in a fissure. The egg is white and
spherical; one which I measured was seven inches
and three-eighths in circumference, and therefore
larger than a hen's egg. The young tortoises, as
soon as they are hatched, fall a prey in great
numbers to the carrion-feeding buzzard. The old
ones seem generally to die from accidents, as from
falling down precipices: at least, several of the
inhabitants told me, that they never found one
dead without some evident cause. . . .

There can be little doubt that this tortoise is
an aboriginal inhabitant of the Galapagos; for it is
found on all, or nearly all, the islands, even on
some of the smaller ones where there is no water;
had it been an imported species, this would hardly
have been the case in a group which has been so
little frequented. . . .

The Amblyrhynchus, a remarkable genus of
lizards, is confined to this archipelago; there are
two species, resembling each other in general
form, one being terrestrial and the other aquatic.
This latter species (A. cristatus) was first
characterized by Mr. Bell, who well foresaw, from
its short, broad head, and strong claws of equal
length, that its habits of life would turn out very
peculiar, and different from those of its nearest ally,
the Iguana. It is extremely common on all the
islands throughout the group, and lives exclusively
on the rocky sea-beaches, being never found, at
least I never saw one, even 10 yards in-shore. It is a
hideous-looking creature, of a dirty black colour,
stupid, and sluggish in its movements. The usual
length of a full-grown one is about a yard, but
there are some even four feet long; a large one
weighed 20 pounds: on the island of Albemarle
they seem to grow to a greater size than elsewhere.
Their tails are flattened sideways, and all four feet
partially webbed. They are occasionally seen some
100 yards from the shore, swimming about; and
Captain Collnett, in his Voyage says, "They go to
sea in herds a-fishing, and sun themselves on the
rocks; and may be called alligators in miniature."

It must not, however, be supposed that they
live on fish. When in the water this lizard swims
with perfect ease and quickness, by a serpentine
movement of its body and flattened tail — the legs
being motionless and closely collapsed on its sides.
A seaman on board sank one, with a heavy weight

attached to it, thinking thus to kill it directly; but
when, an hour afterwards, he drew up the line, it
was quite active. Their limbs and strong claws are
admirably adapted for crawling over the rugged
and fissured masses of lava, which everywhere
form the coast. In such situations, a group of six or
seven of these hideous reptiles may oftentimes be
seen on the black rocks, a few feet above the surf,
basking in the sun with outstretched legs.

I opened the stomachs of several, and found
them largely distended with minced seaweed
(Ulvae), which grows in thin foliaceous expansions
of a bright green or a dull red colour. I do not
recollect having observed this seaweed in any
quantity on the tidal rocks; and I have reason to
believe it grows at the bottom of the sea, at some
little distance from the coast. If such be the case,
the object of these animals occasionally going out
to sea is explained. The stomach contained nothing
but the seaweed. Mr. Baynoe, however, found a
piece of crab in one; but this might have got in
accidentally, in the same manner as I have seen a
caterpillar, in the midst of some lichen, in the
paunch of a tortoise. The intestines were large, as
in other herbivorous animals.

The nature of this lizard's food, as well as
the structure of its tail and feet, and the fact of its
having been seen voluntarily swimming out at sea,
absolutely prove its aquatic habits; yet there is in
this respect one strange anomaly, namely, that
when frightened it will not enter the water. Hence
it is easy to drive these lizards down to any little
point overhanging the sea, where they will sooner
allow a person to catch hold of their tails than jump
into the water. They do not seem to have any
notion of biting; but when frightened they squirt a
drop of fluid from each nostril.

I threw one several times as far as I could,
into a deep pool left by the retiring tide; but it
invariably returned in a direct line to the spot
where I stood. It swam near the bottom, with a
very graceful and rapid movement, and
occasionally aided itself over the uneven ground
with its feet. As soon as it arrived near the edge,
but still being under water, it tried to conceal itself
in the tufts of seaweed, or it entered some crevice.
As soon as it thought the danger was past, it
crawled out on the dry rocks, and shuffled away as
quickly as it could.

I several times caught this same lizard, by
driving it down to a point, and though possessed of
such perfect powers of diving and swimming,
nothing would induce it to enter the water; and as
often as I threw it in, it returned in the manner
above described. Perhaps this singular piece of
apparent stupidity may be accounted for by the
circumstance, that this reptile has no enemy
whatever on shore, whereas at sea it must often fall
a prey to the numerous sharks. Hence, probably,
urged by a fixed and hereditary instinct that the
shore is its place of safety, whatever the emergency
may be, it there takes refuge.

During our visit (in October), I saw
extremely few small individuals of this species, and
none I should think under a year old. From this
circumstance it seems probable that the breeding

season had not then commenced. I asked several
of the inhabitants if they knew where it laid its
eggs: they said that they knew nothing of its
propagation, although well acquainted with the
eggs of the land kind — a fact, considering how very
common this lizard is, not a little extraordinary.

We will now turn to the terrestrial species
(A. demarlii), with a round tail, and toes without
webs. This lizard, instead of being found like the
other on all the islands, is confined to the central
part of the archipelago, namely to Albemarle,
James, Barrington, and Indefatigable islands. To the
southward, in Charles, Hood, and Chatham islands,
and to the northward, in Towers, Bindloes, and
Abingdon, I neither saw nor heard of any. It would
appear as if it had been created in the center of the
archipelago, and thence had been dispersed only
to a certain distance. Some of these lizards inhabit
the high and damp parts of the islands, but they
are much more numerous in the lower and sterile
districts near the coast. I cannot give a more
forcible proof of their numbers, than by stating that
when we were left at James Island, we could not
for some time find a spot free from their burrows
on which to pitch our single tent.

Like their brothers the sea-kind, they are
ugly animals, of a yellowish orange beneath, and of
a brownish red colour above: from their low facial
angle they have a singularly stupid appearance.
They are, perhaps, of a rather less size than the
marine species; but several of them weighed
between 10 and 15 pounds. In their movements
they are lazy and half torpid. When not frightened,
they slowly crawl along with their tails and bellies
dragging on the ground. They often stop, and doze
for a minute or two, with closed eyes and hind legs
spread out on the parched soil.

They inhabit burrows, which they sometimes
make between fragments of lava, but more
generally on level patches of the soft sandstone-
like tuff. The holes do not appear to be very deep,
and they enter the ground at a small angle; so that
when walking over these lizard-warrens, the soil is
constantly giving way, much to the annoyance of
the tired walker. This animal, when making its
burrow, works alternately the opposite sides of its
body. One front leg for a short time scratches up
the soil, and throws it towards the hind foot, which
is well placed so as to heave it beyond the mouth
of the hole. That side of the body being tired, the
other takes up the task, and so on alternately.

I watched one for a long time, till half its
body was buried; I then walked up and pulled it by
the tail; at this it was greatly astonished, and soon
shuffled up to see what was the matter; and then
stared me in the face, as much as to say, "What
made you pull my tail?". . . .

The individuals, and they are the greater
number, which inhabit the lower country, can
scarcely taste a drop of water throughout the year;
but they consume much of the succulent cactus,
the branches of which are occasionally broken off
by the wind. I several times threw a piece to two or
three of them when together; and it was amusing
enough to see them trying to seize and carry it
away in their mouths, like so many hungry dogs

with a bone. They eat very deliberately, but do not
chew their food. The little birds are aware how
harmless these creatures are: I have seen one of
the thick-billed finches picking at one end of a
piece of cactus (which is much relished by all the
animals of the lower region), whilst a lizard was
eating at the other end; and afterwards the little
bird with the utmost indifference hopped on the
back of the reptile. . . .

These two species of Amblyrhynchus agree,
as I have already stated, in their general structure,

and of dimensions comparable only with our
existing whales, swarmed on the land and in the
sea. It is therefore, worthy of his observation, that
this archipelago, instead of possessing a humid
climate and rank vegetation, cannot be considered
otherwise than extremely arid, and, for an
equatorial region, remarkably temperate.

To finish with the zoology: the 15 kinds of
sea-fish which I procured here are all new species;
they belong to 12 genera, all widely distributed,
with the exception of Prionotus, of which the four

Galapagos sheephead wrasse (Cossyphus darwinij. First collected by Darwin while in the Galapagos, and named by
naturalist and friend, Leonard lenyns. This drawing accompanied lenyns's original description. (From ]. E. McCosker and
R. H. Rosenblatt, 1984, Key Environments — Galapagos, Pergamon Press)

and in many of their habits. Neither have that rapid
movement, so characteristic of the genera Lacerta
and Iguana. They are both herbivorous, although
the kind of vegetation on which they feed is so
very different. Mr. Bell has given the name to the
genus from the shortness of the snout; indeed, the
form of the mouth may almost be compared to that
of the tortoise: one is led to suppose that this is an
adaptation to their herbivorous appetites. It is very
interesting thus to find a well-characterized genus,
having its marine and terrestrial species, belonging
to so confined a portion of the world. The aquatic
species is by far the most remarkable, because it is
the only existing lizard which lives on marine
vegetable productions.

As I at first observed, these islands are not so
remarkable for the number of the species of
reptiles, as for that of the individuals; when we
remember the well-beaten paths made by the
thousands of huge tortoises — the many turtles—
the great warrens of the terrestrial
Amblyrhynchus — and the groups of the marine
species basking on the coast-rocks of every
island — we must admit that there is no other
quarter of the world where this Order replaces the
herbivorous mammalia in so extraordinary a
manner. The geologist on hearing this will probably
refer back in his mind to the Secondary epochs,
when lizards, some herbivorous, some carnivorous,

previously known species live on the eastern side
of America.

Of land-shells I collected 16 kinds (and two
marked varieties), of which, with the exception of
one Helix found at Tahiti, all are peculiar to this
archipelago: a single fresh-water shell (Paludina) is
common to Tahiti and Van Diemen's Land. Mr.
Cuming, before our voyage, procured here 90
species of sea-shells, and this does not include
several species not yet specifically examined, of
Trochus, Turbo, Monodonta, and Nassa. He has
been kind enough to give me the following
interesting results: Of the 90 shells, no less than 47
are unknown elsewhere — a wonderful fact,
considering how widely distributed sea-shells
generally are. . . .

I took great pains in collecting the insects,
but excepting Tierra de Fuego, I never saw in this
respect so poor a country. Even in the upper and
damp region I procured very few, excepting some
minute Diptera and Hymenoptera, mostly of
common mundane forms. As before remarked, the
insects, for a tropical region, are of very small size
and dull colours. Of beetles I collected 25 species
(excluding a Dermestes and Corynetes imported,
wherever a ship touches); of these, two belong to
the Harpalidae, two to the Hydrophilidae, nine to
three families of the Heteromera, and the
remaining 12 to as many different families. This

circumstance of insects (and I may add plants),
where few in number, belonging to many different
families, is, I believe, very general. . . .

It was most striking to be surrounded by
new birds, new reptiles, new shells, new insects,
new plants, and yet by innumerable trifling details
of structure, and even by the tones of voice and
plumage of the birds, to have the temperate plains
of Patagonia, or rather the hot dry deserts of
Northern Chile, vividly brought before my eyes.
Why, on these small points of land, which within a
late geological period must have been covered by
the ocean, which are formed by basaltic lava, and
therefore differ in geological character from the
American continent, and which are placed under a
peculiar climate, — why were their aboriginal
inhabitants, associated, I may add, in different
proportions both in kind and number from those
on the continent, and therefore acting on each
other in a different manner — why were they
created on American types of organization? It is
probable that the islands of the Cape de Verd
group resemble, in all their physical conditions, far
more closely the Galapagos Islands, than these
latter physically resemble the coast of America, yet
the aboriginal inhabitants of the two groups are
totally unlike; those of the Cape de Verd Islands
bearing the impress of Africa, as the inhabitants of
the Galapagos Archipelago are stamped with that
of America.

I have not as yet noticed by far the most
remarkable feature in the natural history of this
archipelago; it is, that the different islands to a
considerable extent are inhabited by a different set
of beings. My attention was first called to this fact
by the Vice-Governor, Mr. Lawson, declaring that
the tortoises differed from the different islands, and
that he could with certainty tell from which island
any one was brought. I did not for some time pay
sufficient attention to this statement, and I had
already partially mingled together the collections
from two of the islands. I never dreamed that
islands, about 50 or 60 miles apart, and most of
them in sight of each other, formed of precisely the
same rocks, placed under a quite similar climate,
rising to a nearly equal height, would have been
differently tenanted; but we shall soon see that this
is the case. It is the fate of most voyagers, no
sooner to discover what it is most interesting in any
locality, than they are hurried from it; but I ought,
perhaps, to be thankful that I obtained sufficient
materials to establish this most remarkable fact in
the distribution of organic beings.

The inhabitants, as I have said, state that
they can distinguish the tortoises from the different
islands; and that they differ not only in size, but in
other characters. Captain Porter has described
those from Charles and from the nearest island to
it, namely, Hood Island, as having their shells in
front thick and turned up like a Spanish saddle,
whilst the tortoises from James Island are rounder,

blacker, and have a better taste when cooked. M.
Bibron, moreover, informs me that he has seen
what he considers two distinct species of tortoise
from the Galapagos, but he does not know from
which islands. The specimens that I brought from
three islands were young ones: and probably owing
to this cause neither Mr. Gray nor myself could
find in them any specific differences. I have
remarked that the marine Amblyrhynchus was
larger at Albemarle Island than elsewhere; and M.
Bibron informs me that he has seen two distinct
aquatic species of this genus; so that the different
islands probably have their representative species
or races of the Amblyrhynchus, as well as of the
tortoise. . . .

The distribution of the tenants of this
archipelago would not be nearly so wonderful, if
. . . the different islands were inhabited, not by
representative species of the same genera of plants,
but by totally different genera, as does to a certain
extent hold good: for, to give one instance, a large
berry-bearing tree at James Island has no
representative species in Charles Island. But it is
the circumstance, that several of the islands
possess their own species of the tortoise, mocking-
thrush, finches, and numerous plants, these species
having the same general habits, occupying
analogous situations, and obviously filling the same
place in the natural economy of this archipelago,
that strikes me with wonder. . . .

The only light which I can throw on this
remarkable difference in the inhabitants of the
different islands, is, that the very strong currents of
the sea running in a westerly and W.N.W. direction
must separate, as far as transportal by the sea is
concerned, the southern islands from the northern
ones; and between these northern islands a strong
N.W. current was observed, which must effectually
separate James and Albemarle Islands. As the
archipelago is free to a most remarkable degree
from gales of wind, neither the birds, insects, nor
lighter seeds, would be blown from island to island.
And lastly, the profound depth of the ocean
between the islands, and their apparently recent (in
a geological sense) volcanic origin, render it highly
unlikely that they were ever united; and this,
probably, is a far more important consideration
than any other, with respect to the geographical
distribution of their inhabitants. Reviewing the facts
here given, one is astonished at the amount of
creative force, if such an expression may be used,
displayed on these small, barren, and rocky islands;
and still more so, at its diverse yet analogous action
on points so near each other. I have said that the
Galapagos Archipelago might be called a satellite
attached to America, but it should rather be called
a group of satellites, physically similar, organically
distinct, yet intimately related to each other, and all
related in a marked, though much lesser degree, to
the great American continent. . . .

Darwin and the Galapagos

Three Myths

by Frank J. Sulloway

On 16 September 1835, Charles Darwin landed
in the Galapagos Islands and began five weeks of
collecting and observing in this famous "laboratory
of evolution." While in the Galapagos, the 26-year-
old Darwin visited four of the major islands, and,
from the H.M.S. Beagle, he glimpsed numerous
others. Altogether he spent 19 days on land in the
Galapagos -- five days on Chatham; four on
Charles, where he visited the highlands settlement;
one day at Tagus Cove on Albemarle Island; and
nine days on James, where he collected extensively
and spent three days in the highlands (Figure 1).

By current research standards, Darwin's
Galapagos visit was remarkably brief. And yet his
encounter with these islands was seemingly
decisive for his biological thinking. As he wrote in
the second edition of his journal of Researches:

The archipelago is a little world within itself, or
rather a satellite attached to America, whence it
has derived a few stray colonists, and has
received the general character of its indigenous
productions. Considering the small size of these
islands, we feel all the more astonished at the
number of their aboriginal beings, and at their
confined range. Seeing every height crowned
with its crater, and the boundaries of most of
the lava-streams still distinct, we are led to
believe that within a period geologically recent
the unbroken ocean was here spread out.
Hence both in space and time, we seem to be
brought somewhat near to that great fact -
that mystery of mysteries -- the first appearance
of the new beings on this earth. (1845: 377-78)

When and how Darwin solved this great
"mystery of mysteries," and particularly the role his
Galapagos visit played in this regard, have become
the subject of a considerable legend in the history
of science.

According to the legend, Darwin's
Galapagos visit first provided him with irrefutable
evidence for the mutability of species and
converted him, eureka-like, to the theory of
evolution. Actually, the impact of the Galapagos
was largely retrospective. Darwin was first alerted
to the evolutionary significance of the Galapagos
species by the vice-governor, Nicholas Lawson,
who informed him that he could tell "with
certainty" from which island any tortoise had been
brought. Darwin was on Charles Island at the time;
and according to David Lack, among other
commentators, he was sufficiently impressed to
begin separating his collections of finches and
other species by island, thus securing the necessary
biological evidence to back up the vice-governor's
extraordinary claim. What Lack and others did not
appreciate, however, was that the bulk of the
locality information on Darwin's type specimens
and in his postvoyage publications was actually
derived, after the voyage, from the carefully
labelled collections of three other Beag/e shipmates
(all naval personnel). Why Darwin initially failed to
heed the vice-governor's remarks about the
tortoises must be understood in terms of the
intimate relationship between a received theory
like creationism, no matter how erroneous, and the
gathering and interpretation of scientific evidence.

To begin with, it would never have occurred
to a creationist, which Darwin still was in 1835, to
label his collections according to island of origin
within a small archipelago. As part of a presumed
"center of creation," the Galapagos would have
been expected to exhibit a uniform flora and fauna
by island, making such detailed locality
designations superfluous. In this regard, it is
noteworthy that those Beagle specimens that were
carefully labelled by island were collected by the
nonscientists on board, who presumably did not
realize how unnecessary such information really
ought to have been.

We also tail to appreciate how complex and
confusing the Galapagos evidence must initially
have been, especially to a nonspecialist and

nonsystematist like Darwin. It is not just the theory
of evolution that introduces unifying order into
many of the enigmas of Galapagos biology;
creationism also made a certain reasonable sense
out of the facts. From his specimen notebooks and
manuscript notes it is clear, for example, that
Darwin mistook many species of "Darwin's finches"
for the forms that they, through adaptive
evolutionary radiation, now appear to mimic. Thus
he thought the warbler finch was a "Wren"; and he
described the large-beaked ground finch as a
"Grosbeak" and the cactus finch as an "Icterus"
the genus to which belong the orioles, blackbirds,
and certain other forms possessing a long pointed
bill. It is perhaps not surprising then that Darwin,
having failed to recognize the closely related
nature of the Galapagos finches, also failed to
suspect that their island distributions might vary
within the archipelago.

The evolutionary evidence provided by the
famous Galapagos tortoises was also similarly
clouded at the time of Darwin's visit. This taxon
was then believed by most naturalists to have
originated in the islands of the Indian Ocean -
hence its erroneous name Jestudo indicus -- and
to have been transported to the Galapagos by
buccaneers. Thus when Darwin was informed that
the tortoises differed by island, he probably initially
thought it was a matter of local variations somehow
induced by transportal to a new and unnatural
environment. Moreover, those tortoises actually
seen by Darwin, on Chatham and James, were too
similar to be distinguished "with certainty"; so the
evidence was not as striking, from Darwin's
personal observations, as the vice-governor had
claimed.

In any event, since tortoises were not
supposed to be native to the Galapagos, such
differences did not apparently bear directly on the
question of what was uniquely "Galapagean," if
anything, about the Galapagos. So little value did
Darwin place upon the tortoise evidence that he
not only failed, at the time of his visit, to collect
specimens for scientific purposes, but he
apparently joined his Beagle shipmates in eating
the last of some 30 large tortoises during the cruise
to Tahiti. It was only a decade later that Darwin
finally encountered Captain David Porter's (1815)
description of the dome-shaped and saddleback
forms of tortoise and was able to insert this
information into the second edition of his journal of
Researches (1845: 394).

The Origin of Species (1859) was never in
any real danger, however, of being sacrificed for a
bowl of tortoise soup. Darwin had noticed, while
still in the Galapagos, that the mockingbirds
differed by island; and he had taken care to
separate these specimens from the four islands he
had visited. Approximately eight months after
leaving the Galapagos he returned to this problem
in his "Ornithology" notes. There he compared this
anomalous finding to that previously reported to
him about the tortoises. Although he was still
inclined to suspect that his mockingbirds were
"only varieties" rather than true species, he
nevertheless speculated that "If there is the

slightest foundation for these remarks the zoology
of Archipelagoes -- will be well worth examining;
for such facts [would inserted] undermine the
stability of Species" (1963 [1836]: 262). Darwin
thus began, in a tentative but probing manner, the
real process of "discovery" about the Galapagos -
a process that lay not so much in his observations
or collections during his brief visit, but rather in his
various reconsiderations of this evidence after his
departure.

Following his return to England in the
autumn of 1836, Darwin had many opportunities to
re-evaluate the Galapagos evidence as expert
systematists began to work out his voyage
collections and he prepared his lournal of
Researches for publication. In early March of 1837,
he met with the celebrated ornithologist John
Gould to discuss the results of Gould's examination
of his voyage birds. Gould had immediately
appreciated the anomalous but closely related
nature of Darwin's Galapagos finches, including the
warbler finch, and had named 13 species in three
subgenera. In addition, Gould had pronounced as
distinct three of the four island forms of Darwin's
Galapagos mockingbirds, thus confirming the
suspicions Darwin had previously felt might
"undermine the stability of Species." Perhaps just
as importantly, Gould convinced Darwin of the
highly endemic character of the Galapagos
ornithology as a whole, something that Darwin,
who had not had access to museum collections
during the voyage, had not previously realized.
These taxonomic opinions, together with a number
of others relating to his collections from the South
American continent, finally convinced Darwin that
species were indeed mutable and sparked his
decision to begin collecting facts that might bear
on this question. He subsequently commented in
this connection: "In July [1837] opened first
notebook on 'Transmutation of Species' - - Had
been greatly struck from about Month of previous
March on character of S. American fossils -- and
species on Galapagos Archipelago. These facts
origin (especially latter) of all my views."

In the wake of his conversion to the theory
of evolution, Darwin quickly realized his voyage
oversight in failing to label his Galapagos
specimens by island. He therefore set out to rectify
this problem as best he could by asking other
Beagle shipmates, including Captain Robert
FitzRoy, to supply him with the missing evidence.
Unfortunately, later curators at the British Museum
failed to appreciate that Darwin's published locality
designations in the Zoology of the Voyage of H.M.S.
Beagle (1841) were not derived from his own
collections; and where such information was
missing from his own type specimens, they added
it to some of the labels, creating a number of
erroneous localities. Darwin, moreover,
compounded the problem by guessing where eight
of his own finch specimens had come from; and in
several instances he clearly guessed incorrectly.
These various confusions over the type specimen
localities created a taxonomic nightmare for
subsequent ornithologists, who naturally puzzled
over the conflicting and aberrant locality

Figure 1 . Darwin's route
through the Galapagos in
H.M.S. Beagle. He visited the
four shaded islands and made
several inland excursions, also
indicated on the map. The
occasionally zigzag nature of
the Beagle's route reflects the
vagaries of winds and currents
in the age of sail.

CHATHAM r^.

designations on Darwin's specimens and found
themselves hard pressed to reconcile this
information with present-day distributions of
Darwin's finches.

Fortunately, clarification of the retrospective
and borrowed nature of the localities on many of
Darwin's type specimens has now resolved most of
these problems, including the status of several
long-debated forms of Darwin's finches. In
particular, Geosp/za magnirostris magnirostris, an
extinct form of the large-beaked ground finch, was
collected by FitzRoy and others on Chatham and
Charles islands, where David Steadman (1981,
1984) has recently found fossil evidence of this
subspecies. Similarly, both Darwin and FitzRoy
collected specimens of another extinct subspecies
on Charles Island --a particularly large-billed form
of the sharp-beaked ground finch ("C. nebulosa"
Gould).

Although Darwin (1845: 395) later suggested,
based on the joint Beagle collections, that the
Galapagos finches might have different geographic

distributions, he was also aware that the case was a
complex one and that his own data on the subject
were meagre and probably suspect. Partly for this
reason he did not mention his celebrated
Galapagos finches in the Origin of Species (1859). It
is only in this century, after the splendid
ornithological studies of Harry Swarth (1931), David
Lack (1945, 1947), and many other researchers,
that these finches have become such a convincing
paradigm of evolution in action. In keeping with
the Darwin-Galapagos legend, however, much of
this modern evidence is often erroneously
attributed to Darwin. For example, he never saw all
13 species of Galapagos finches (Gould's 13
"species" encompassed only nine of the presently
recognized forms), and he was also unaware that
differences in the beaks were correlated with
differences in diets.

Even after he had finally become an
evolutionist in 1837, Darwin's understanding of the
Galapagos Islands continued to undergo a slow
evolution of its own. The mockingbirds and

tortoises had convinced him of the importance of
geographic isolation in the evolution of new
species; and in 1838, after reading Malthus's Essay
on the Principle of Population (1798), he hit on the
theory of natural selection. (Even this important
insight, however, was not as sudden as Darwin
later recalled.) For approximately a decade more
he nevertheless failed to understand why evolution
should promote widely divergent species on
islands, like the Galapagos, that are seemingly
identical in climate and general geographic
character.

Darwin solved this vexing problem only in
the mid-1 840s after reading Joseph Hooker's
reports on the flora of the Galapagos. Hooker had
found that numerous representative species were
indeed present on the separate islands, as Darwin
had always suspected but had never been able to
prove conclusively. In July of 1845, Darwin wrote
to his friend: "I cannot tell you how delighted and
astonished I am at the results of your examination;
how wonderfully they support my assertion on the
differences in the animals of the different islands,
about which I have always been fearful."

Darwin was equally impressed with
Hooker's (1847) discovery that the different islands
possessed plants that were apparently random
colonists, present only on one island. In the margin
of his copy of Hooker's paper Darwin wrote: "so
the flora of different isldfs] must be very different
independently of representation." Darwin now
began to appreciate that although the various

islands in the Galapagos might look superficially
similar, they were biotically quite distinct. These
biotic differences, moreover, must provide natural
selection with a wide scope for expression, thus
explaining how representative species had evolved
so easily on each island. This basic idea, which
Darwin developed in the 1850s into his principle of
divergence, altered much of his general thinking
about evolution and was given a prominent place
jn the Origin of Species (1859). Thus Darwin
required almost two full decades to understand the
biological significance of his Galapagos findings and
to integrate them into his theory of evolution by
natural selection.

The Darwin-Galapagos Legend

The publication of the Origin of Species not only
revolutionized the biological sciences, but it also
made Darwin into a celebrated intellectual hero — a
man thoroughly worthy of scientific deification and
hence destined to become the subject of legend.
And because myths and legends, above all else,
gravitate toward the problem of origins, Darwin's
discoveries increasingly became enshrouded by the
typical misconceptions of reconstructed "heroic"
history. Accordingly, the true story of Darwin's
conversion to the theory of evolution is a far cry
from the Darwin-Galapagos legend that has arisen
in the wake of Darwin's scientific triumph, and that
adorns so many of the biology textbooks today. In
fact, the legend, which is composed of three major
component myths, tends to obscure precisely what

Darwin's Calapagos
mockingbird specimens (British
Museum of Natural History,
Tring). From top to bottom (in
the order that Darwin collected
them): the Chatham Island
mockingbird (Nesomimus
melanotis), the Charles Island
mockingbird^, trifasciatus),
and the Galapagos
mockingbird (N. parvulus).
Darwin collected two
specimens of the latter, one on
Albermarle and the other on
lames islands. The fact that
Darwin procured only four
specimens during his
Calapagos visit — one from
each island — shows that he
was collecting within a
creationist perspective. To an
evolutionist there can be no
single "type " specimen, since
the variation within the species
is an important part of its
genetic nature and not simply a
"deviation from the type." (All
photographs are by the author)

Three subspecies of Galapagos tortoise. Left: a pair of
Chatham Island tortoises (Geochelone elephantopus
chathamensis), displaying relatively dome-shaped carapaces.
Right: the Hood Island tortoise (G. e. hoodensis), an extreme
saddleback form similar to the now-extinct Charles Island
race (G. e. galapagoensis). Below: the lames Island tortoise:
(G. e. darwinii), a dome-shaped form. Darwin unfortunately
saw only the two similar dome-shaped forms.

it pretends to explain, namely, the nature of
scientific insight.

The first of these component myths is that of
Darwin's "eureka-like" conversion during his brief
visit to the Galapagos Islands. It may appeal to our
romantic conception of scientific discovery to
imagine the lone voyager suddenly throwing off the
shackles of creationist thinking when finally
confronted, in the Galapagos, with a microcosmic
paradigm of evolution in action. But this myth, for
all of its inherent allure, is both wrong and
misleading. What this myth especially tends to
obscure is the fascinating question 'Why Darwin?'
That is to say, why was it that Darwin, and no one
else, was converted by evidence that was widely
known to many other contemporary naturalists-
naturalists who, like Richard Owen and John
Gould, were often far superior to Darwin in their
experience and abilities as systematists? The answer
to this question is closely associated with the real
nature of Darwin's genius as a scientist. As the far-
seeing amateur among specialists, Darwin exhibited
his unique intellectual caliber in the pattern of

"gifted individualism" that manifested itself in the
process of his conversion. While other naturalists
stood by and calmly rationalized the Galapagos
evidence in creationist terms, Darwin — virtually

alone — took up the heterodox challenge offered
by that evidence. Expressed another way, the
Galapagos did not make Darwin; if anything,
Darwin, through his superior abilities as a thinker
and a theoretician, made the Galapagos; and, in
doing so, he elevated these islands to the
legendary status they have today.

The second of the three component myths
associated with Darwin and the Galapagos is the
myth that these islands provided him, at an early
stage, with a basic paradigm for his theory of
evolution by geographic isolation and natural
selection. As I have shown in the case of Darwin's
finches, nothing could be further from the truth;
and the same conclusion applies to Darwin's
Galapagos observations as a whole, which were
only slowly incorporated into his final theory. Thus
the Origin of Species was ultimately the product of
24 years of thinking and further research (1835-
59), not the five weeks that Darwin spent in the
Galapagos Islands or even the five years that he
spent accompanying H.M.S. Beagle around the
world. True, the Galapagos certainly provided
Darwin with some crucial hints; but Darwin's full
understanding of both evolution and the Galapagos

The remarkable diversity in the forms of the Galapagos
finches is shown here by three species that initially misled
Darwin into thinking they were members of separate families
or subfamilies: the large-beaked ground finch (Ceospiza
magnirostris), using its powerful jaws to crush a large seed;
the cactus finch (G. scandens), feeding on the flowers of
Opuntia; and the diminutive warbler finch (Certhidea
olivacea) looking for insects in the highland Scalesia forests.

case required almost as long as it took him to
publish the Origin of Species.

Moreover, much of Darwin's evolutionary
argument, as finally presented in the Origin, had to
be constructed from alternative sources, owing to
Darwin's failure to appreciate, and to collect, the
necessary Galapagos evidence in 1835. Other
scientists have been collecting that "necessary"
Galapagos evidence ever since, which leads me to
the third of the three component myths
encompassing the Darwin-Galapagos legend.

This third and last myth involves the notion
that Darwin singlehandedly discovered almost
everything there is to know about evolution in the
Galapagos — or at least everything of basic
importance — and hence that subsequent research
in these islands has merely been a sort of mopping-
up operation characteristic of "normal,"
postrevolutionary science. This myth, promulgated
in the biology textbooks and especially in the
popular literature about Darwin and the Galapagos,
is largely a natural extension of the first two
Darwin-Galapagos myths.

As a typical manifestation of this third myth,
Darwin is frequently credited with insights about
his famous Galapagos finches that were actually
the product of extensive post-Darwinian
ornithological research. For example, in spite of
Darwin's own famous journal (1845: 380) remark
about one species of finch appearing to have been
"modified for different ends," Darwin was by no
means personally convinced that all 13 species of
Galapagos finches (especially the warbler finch)
were indeed derived from a single ancestor (see
also Darwin, 1841: 105). Darwin's lingering doubts

about the finches' possible common ancestry
apparently contributed to his decision, when
writing the Origin of Species, to omit any specific
reference to this now famous biological paradigm
of "evolution in action." During the remainder of
the 19th century, ornithologists generally believed
Darwin's finches were descended from two or
three different ancestors — a warbler, a ground
finch, and a separate form that gave rise to the six
species of Camarhynchus. This issue of ancestry
was not resolved for more than half a century after
the Origin of Species was published.

David Lack's classic book Darwin's Finches
(1947) did much to perpetuate this third aspect of
the legend, even though Lack himself personally
knew better. Indeed, Lack, in reversing his original
position on the possible adaptive significance of
the beaks among the different species of Darwin's
finches (1945, 1947), went through much the same
experience of ex post facto 'discovery' that Darwin
himself did. For it was only after leaving the
Galapagos Islands that Lack reached his new
theoretical position and then realized the need for
the kind of follow-up studies of the finches'
feeding behavior that various other ornithologists
have subsequently carried out.

Similar "delayed discoveries" have
undoubtedly characterized the work of numerous
other Galapagos researchers. Unlike Darwin,
however, they have often had the opportunity to
return to the Galapagos Islands in order to collect
crucial data, and to make observations, that
previously seemed unimportant. Thus the history of
research in the Galapagos Islands has been
anything but the history of "mopping up" the
scientific tidbits that Darwin left behind. Rather, it
is only after repeated expeditions by six
generations of post-Darwinian scientists that the
Galapagos archipelago has yielded — with a
seeming air of reluctance — many of its richest
biological treasures to the world of science. And
even today, after so much scientific progress,
almost as many questions remain about evolution
in the Galapagos as there are answers to the
mysteries that Darwin and others have successfully
resolved.

Of all the scientists who have made
important discoveries in the Galapagos, only to
realize later that they have merely scratched the
scientific surface and thereby created the need for

Darwin in 1840, at age 3 1, five years after the Beagle voyage.
(By George Richmond, courtesy of Downe House, Downe,
England)

further research, Charles Darwin perhaps
expressed it best. In 1846, shortly after Joseph
Hooker had so delighted him with the results of his
analysis of Darwin's Galapagos plants, Darwin
declared to his friend: "The Galapagos seems a
perennial source of new things." The Darwin-
Galapagos legend notwithstanding, these famous
islands will doubtless remain "a perennial source of
new things" in science; and no one would be more
disappointed than Darwin if this were not the case.

Frank /. Sulloway is a MacArthur Fellow in the Department
of Psychology and Social Relations, Harvard University,
Cambridge, Massachusetts.

Selected Readings

Darwin, C. R. 1839. Journal of Researches into the Geology and

Natural History of the Various Countries Visited by H.M.S.

Beagle under the Command of Captain FitzRoy, R.N. from

1832 to 1836. London: Henry Colburn
Darwin, C. R., Ed. 1841 . The Zoology of the Voyage of H.M.S.

Beagle, under the Command of Captain FitzRoy, R.N., during

the Years 1832-1836. Part III: Birds. London: Smith, Elder &

Co.
Darwin, C. R. 1845. Journal of Researches into the Natural History

and Geology of the Countries Visited during the Voyage of

H.M.S. Beagle Round the World, under the Command of

Capt. FitzRoy, R.N. 2nd ed. London: John Murray.
Darwin, C. R. 1859. On the Origin of Species by means of Natural

Selection, or, The Preservation of Favoured Races in the

Struggle for Life. London: John Murray.
Darwin, C. R. 1887. The Life and Letters of Charles Darwin,

Including an Autobiographical Chapter. Edited by F. Darwin. 3

vols. London: John Murray.
Darwin, C. R. 1958, 1876. Autobiography: With Original

Omissions Restored. Edited with Appendix and Notes by his

grand-daughter. N. Barlow. London: Collins.
Sulloway, F. J. 1982. Darwin's conversion: The Beagle voyage and

its aftermath, lournal of the History of Biology 1 5: 325-396.

Whalers, Whales,
and Tortoises

The bark Morning Star of New Bedford: at Albemarle Island, Galapagos, July 27 to August 5, 1858; at Chatham Island from June
27 to luly 1 1, 1861. Total catch of tortoises, 212. (From C.H. Townsend, 1927)

by Bruce C. Epler

I or most of us, mention of the Galapagos brings to
mind images of tortoises, volcanoes, marine iguanas,
or blue-footed boobies. But, it was the resources
hidden beneath the seas surrounding the
archipelago, namely whales, that brought visitors by
the thousands between 1790 and the early 1900s.
They came in search of sperm whales, and
sometimes seals. What is less well-known is that they
left the islands with large numbers of tortoises stored
in their holds. The impact of their activities lingers
on.

/ hove to and sent the chief mate on shore to
sound and land. At eight, P.M. he returned with
green turtle and tortoises (galapagos), turtle doves
and guanas but they saw no esculent vegetable,
nor found any water that was sufficiently
palatable to drink.

— Captain Colnett aboard the British whaler
Rattler, June 24, 1793.

G. W. Shuster, writing in a 1983 International
Whaling Commission report, recounts the Rattler's
return visit in April, 1794: "They saw many
spermacet(i) whales, especially young ones. They
killed five here and (Captain) Colnett believed he
had discovered the general rendezvous of these
whales from the coast of Mexico, Peru, and the Gulf
of Panama who came here to calf."

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f/gure 7. The South Pac/ftc, as seen by Samuel Enderby, who outfitted the British ship Rattler, (he first whaler to visit
the Galapagos. This dearly illustrates the role of the Galapagos as the whaler's gateway to the Pacific. (Source: I. T.
Sanderson, 1956)

The (op portion of a page from the logbook of the Bark Chili, captained by Benjamin 5. C/ar/c. The sh/p visited the Galapagos
several times during the mid-1 850s, and returned to New Bedford on lune 8, 1960, with 704 barrels of sperm oil and 128 barrels
of whale oil. (Photo courtesy of the Kendall Whaling Museum)

Whaling in The Pacific

In 1788, the British Ship Amelia was the first vessel to
round Cape Horn and enter the Pacific in search of
sperm whales. Americans were quick to follow, as no
less than six American vessels rounded the Cape in
1791. Commercial mariners had never taken to the
seas in the same magnitude as the whalers who
came to the Pacific, in large numbers, from the
United States, Britain, France and, in smaller
numbers, from Holland, Spain, and the German
states of Hamburg and Bremen.

The British ship William, the second whaler to
enter Galapagos waters (the Rattler being the first),
caught 42 of its 100 sperm whales around the islands
in an 18-day period in January 1797, and the British
whaleship Cyrus shortly afterwards achieved a full
load in only a year and a half of cruising around the
Galapagos.

From this modest beginning, the reputation of
the rich whaling ground and unique inhabitants to
be found on and around the Galapagos Islands
spread through the world's whaling fleets. During
the next century, tens of thousands of whales, fur
seals, and tortoises would be taken. Buccaneers and
explorers of the 17th century, including Captains
Dampier, Davis, Cook, Wafer, Knight, Cowley, and
Eaton, who had earlier used the islands as a retreat
from which to raid and burn coastal towns and loot
Spanish ships, consumed a goodly number of
tortoises, but the amount was minuscule in
comparison to the demands of the whalers.

Once in the Pacific, the whalers pressed their
search along the entire coast of South America. Rich
new whale grounds were discovered as the
whaleships, carried by the currents and prevailing
winds, crossed from the Galapagos along the
Equator, where sperm whales congregated in large

numbers, to the Gilbert (Kiribati) and Ellice Islands
(Tuvalu). From these so-called "on the line" grounds,
they sailed southward to the Vasquez grounds, and
eventually to New Zealand; and north to Japan
grounds (Figure 1).

New England Whaling in The Galapagos

R. Langdon (see Selected References) lists the Lady
Adams of Nantucket, which arrived at the islands in
May, 1803, as the first documented American
whaling vessel to reach the Galapagos. The islands
soon became a favorite cruising grounds for
American whalers. In one instance, for example, in
May 1809, a ship off Narborough (Fernandina) Island
"spoke" 10 other whaleships in the space of one
day. Melville, during a stop in the islands on board
the Acushnet in 1841, writes,

The day after we took fish at the base of this
Round Tower (Redondo Rock), we had a fine
wind, and shooting round the north headland,
suddenly decried a fleet of full thirty sail, all
beating to the windward like a squadron in line.
A brave sight as ever man saw. A most
harmonious concord of rushing keels. Their thirty
kelsons hummed like thirty harp-strings, and
looked as straight whilst they left their parallel
traces on the sea. But there proved too many
hunters for the game. The fleet broke up, and
went their separate ways out of sight, leaving my
own ship and two trim gentlemen of London.

The reasons why whalers congregated in the
Galapagos are easily understood: the surrounding
grounds were rich in whales and the ideal jumping-
off point for trips into the Pacific. Water, wood, fresh
provisions, good fishing, sand (for the cooper or for

use in scrubbing the deck), dry salt (in uplifted
craters), and sheltered harbors were available. They
also were ideally situated to come and go from the
"Offshore ground" to the southwest, the largest
ground in the Eastern Pacific. The presence of large
numbers of "succulent" tortoises provided an
additional attraction.

A contributing factor in increasing the
popularity of the archipelago was refusal by the
Spaniards to recognize American sovereignty during
the Revolutionary War and the War of 1 81 2.
American vessels in Spanish ports along the South
American coasts were seized; those at sea were
captured either by British or by Chilean and
Peruvian pirates emboldened by wartime conditions.
During both wars, prudent captains, in increasing
numbers, sought the refuge afforded by the
Galapagos. The islands became a well known
rendezvous for New England whalers.

Most American logs are filled with reports of
"gamming," getting together, or "companioning,"
cruising with vessels, often from their home port,
while in the Galapagos. Vessel traffic was such that
some time prior to 1 793, a post office, constructed
of a large covered tortoise shell, and identified by a
black sign as "Hathaway's Post Office," was
established on Floreana Island to facilitate mail
delivery between passing ships and their countries of
origin. (This post office and service are still in
operation today.

The Taking of Tortoises

Tortoises, because of their ability to live confined
within the holds of ships, with little or no food and
water, for extended periods of time (more than a
year by some accounts) were prized by whalers as
an important source of fresh meat during long
passages at sea. Admiral Porter, 1813, after bringing
"fourteen tons" from Santiago on board the Tartar
mentions, "They require no provisions or water for a
year.. . . They have been piled away among the
casks in the hold of a ship, where they have been
kept for eighteen months."

Searching the islands for tortoises is, generally,
an ordeal. In many areas, walking involves balancing
on jumbled masses of broken lava which crumble
and slip under foot, frequently penetrating shoe
leather, and circumventing crevices while
maneuvering through tangled thickets of brush and
cactus under the weight of an unrelenting tropical
sun. More than one man was lost in the process.

Sept. 25, 1842
Daylight Sept. 26
Sept. 27

two boats came with 22
(tortoises) lost one man

sent all hands to look for
lost man

could not find him, left
bread and water and
directions in a bottle, if
anyone should ever find
him.

Log of the ship Chili at Isabela

Even though these creatures weigh up to 550
pounds, it is logical to presume that tortoises

ranging from about 25 to 75 pounds were
preferred, as they are more easily carried and,
likely, the most tender. A tortoise per man was the
usual load, the carrying of which was called
"backing them down." Straps or belts were used to
facilitate carrying. Larger tortoises were attached
to oars so that they could be carried by two or
more men. Others were killed and had their meat
removed on the spot. In some instances, they
were rolled onto their backs and dragged by ropes
attached to their legs. (The difficulty in
transporting these animals is one explanation why
populations on smaller islands with low elevations
fared poorly in comparison to populations in the
distant highlands of larger islands.

Referring to tortoises, Melville writes:

. . . most mariners have long cherished a
superstition, not more frightful than grotesque.
They earnestly believe that all wicked sea-officers,
more especially commodores and captains, are at
death (and in some cases, before death)
transformed into tortoises; thenceforth dwelling
upon these hot aridities, sole solitary lords of
Asphaltum.

Loading tortoises aboard ship was likewise no
simple process. One account of the ships Coral and
Hope at San Cristobal Island between February 27
and August 2, 1948 reads:

This [the loading] was very hard and sometimes
very dangerous work, as we were obliged to send
them down to the boat by means of a rope from
the tops of the cliffs which were perpendicular
and above a hundred feet above the surface of
the seas. Our method was this, we went in with
two boats as near as possible, for we could not
go in close under the cliffs on account of the surf
running so very high. We anchored one boat and
then made a rope fast to her and all the men got
into the other boat taking with them the end of
the rope by which we could slack the boat or in
haul her out at pleasure.

[Once a continuous loop of line from the cliff to
the boat was arranged] the terrapin were then
made fast to one part one at a time and lowered
about half way down when another was made
fast and lowered away, and so we were kept
going, the line going round and round so that
one part was going up when the other was going
down and 3 terrapin on it all the time. When we
got one boat full she was taken to the ship and
discharged and the line was anchored by means
of a stone. In this manner we got off about 200 of
them. Two or three of them fell whilst lowering
them down and were dashed to pieces on the
rocks.

Quantities of Tortoises Harvested

It is impossible to estimate the number of tortoises
carried away in the holds of whaling vessels that
frequented the islands for more than a century.

Commercial

In addition to forto/ses, large numbers of fur
seals, along with some sea lions, were harvested
in the islands. The first mention of the
commercial potential for harvesting these animals
is found in the log of the British ship Rattler
which reads:

We saw but few seals on the beach, either
of the hairy [sea lions] or furry [fur seals]
species. This circumstance, however, might
be occasioned, by its not being the season
for whelping; as those which were killed by
us, had some time to go with young; but a
few hundred of them, might at any time be
collected without difficulty, and form, no
inconsiderable addition, to the profits of a
voyage.

A review of logs at the Kendall Whaling
Museum, Sharon, Massachusetts, reveals that
whalers were well aware of their value, with
entries such as; "only found 3 Turipin got 4 hair
seals," or "boat returned with wood and two seal
skins," are common.

Whalers were not the only American
commercial vessels to harvest animals on the
islands. Sealers frequented the islands, in much
reduced number compared to whalers,
throughout the 18th century. A partial record of
documented sealing activities is presented below.

7876: During Fanning' s voyage, in 1816, 8,000
fur seals and 2,000 sea lions were taken at
the Galapagos Islands.

1823: Benjamin Morrell, "We remained among
these islands about two months during
which period we took about 5,000 fur seal
skins."

1825: Morrell, on a return voyage found only a
few seals that were taken at the south end
of Isabela Island. However, when the
volcanic island unexpectedly erupted on
February 14, 1825, Morrell' s ship was
anchored off Fernandina. His account
reads:

Our ears were suddenly assailed by a
sound that could only be equalled by ten
thousand thunders bursting upon the air
at once; while, the whole hemisphere was
lighted up with a horrid glare that might
have appalled the stoutest heart.
... At the time the mercury in the
thermometer was at 147, but on
immersing it into the water, it instantly
rose to 150. Had the winds deserted us
here, the consequences must have been
horrible.

1872-1880: Capt. C. W. Reed made four
sealing voyages to the Galapagos Islands
between 1872 and 1880, during which
about 6,000 seals were taken. The skins
are said to have been less valuable than
those from Guadalupe, Santa Rosa, and
Santa Cruz Islands.

1897: Captain W. P. Noyes, of the schooner
Prosper, of San Francisco, visited the
Islands in 1897, and between luly 16 and
October 19 secured 224 seals, 139 of
which were females. Although the
logbook records of this voyage state that
the seals were procured at distances
varying from 1/2 to 7 miles from shore, the
master subsequently stated that some of
the animals were killed in caves and
elsewhere on land.

During its peak, the American whaling fleet alone
contained more than 700 vessels that made repeated
voyages into the Pacific. Information contained in
foreign logs is not included here and is a matter of
conjecture.

Thousands of log books have, regretfully,
been lost over time. The ones that exist are working
documents containing information of concern to the
vessel owner, and provide only sporadic details on
activities on the islands and places visited. We know
that many vessels that took tortoises failed to record
it. Given the demand for fresh food on board, one
may presume that each vessel at least searched for
tortoises and most were successful.

An additional problem is that many entries
read "took 24 plus many," "employed giting turpin,"*
"employed Turpining," "stowing Terrapin," or "seven

boat loads more," so it is impossible to assign values.
Even the number of animals required to fill a boat is
undefined as the sizes of the boats and tortoises
varied over the years. The log of the bark Morning
Star at San Cristobal on July 10, 1861 reads, "all three
boats came on board each one brought 20 Turpin."
Captain Barnard's narrative of the ship Millwood says
"Mr. Coles had forty-five terrapin in the boat."

The most comprehensive work assessing the
impact that American whalers had on the Galapagos
tortoise population is a 1923 study by C. H.
Townsend. It is based on log books from 79 whaling
vessels that made 189 visits to the islands between
1 83 1 and 1 868 for the purpose of securing tortoises.

* Various spellings of the word terrapin are used by the
whalers when referring to tortoise.

Sealing

The Galapagos sea lion. (Photo by D. /. H. Phillips)

Records reviewed by the author indicate
that 22,485 fur seals and 2,000 sea lions were
killed, yet this represents only a small proportion
of the number actually taken.

By the 7890s Galapagos fur seals were
thought to have been exterminated. The
Albatross during its 1891 voyage to the islands
"found that a scattered remnant of a herd still
frequented the more inaccessible rocks of the
archipelago. " Subsequent sealing voyages
"resulted in the killing of all seals that could be
found." An 1899 government report states that "It
was a matter of surprise to those interested in the
subject that during the past month (December,
1897) a vessel arrived at San Francisco from the
Galapagos with a catch of 224 seals, as no one
had anticipated that the race had survived."

A California Academy of Sciences
expedition (1905-06) visited the islands to
conduct the first comprehensive survey of the
archipelago. During 1 2 months of investigation,
only one fur seal was observed.

The present population of seals is
estimated to be between 30,000 and 40,000.
Small clusters are found throughout the islands
with the majority of the population located in the
northwest of the archipelago on Marchena, Pinta,
Isabela, and Fernandina islands. Sea lions, whose
low-valued pelts were not sought by sealers, are
widespread throughout the islands. Their
population is between 20,000 and 50,000 but the
sea lions have been subject to an epidemic of
seal pox, and periodic influences of the El Nino,
both of which are reducing their numbers.

— BCE

A summary of the total number of tortoises
Townsend estimated as taken by ships in his sample
is presented in Table 1 . The largest catches of
tortoises over specified periods of time recorded by
Townsend from individual islands are:

1834 ship Moss Floreana Island 350 9 days

1831 ship Isabella Espanola Island 335 5 days

1831 ship Hesper Espanola Island 250 6 days

1837 ship Omega San Cristobal 240 9 days

Island

In one instance, the log of the ship Uncas
from Woods Hole contains the following entries:

13th March, 1834

Came to anchor at James
[Santiago] all hands
employed after Tirripen.

14th- 17th March
18th March

All hands employed after

Turpin.
At 4 P.M. got under way and

steered N.N.W. with 416

terrpins.

Table 1. Total catch of tortoises arranged by decades.

Number of
Decades Tortoises Average/Visit visits

1831-39

4853

1840-49

4379

1850-59

2334

1860-68

1447

Totals

13,013

189

It is worth noting that the totals and average numbers of tortoises
taken per visit steadily declined, and that a similar trend is evident in
average number of visits each decade — all indicating a rapidly shrink-
ing population.

Table 2. Total catch of tortoises (1831-1861)1, number of races per island, status in 19062, and present estimated population3.

Number of Number
Island races4 Taken

Status in 1906

Status in 1974

San Cristobal

4,798

Nearly Extinct

500-700 discovered in 1972

Floreana

1,775

Extinct

Espanola

1,698

Very rare

20 to 30*

Santiago

1,048

Rare

500 to 700s 6

Pinta

455

Rare

One tortoise found in December, 1971

Pinzon

356

Fairly abundant

150 to 200s

Santa Cruz

366

Not rare

2,050 to 3, 100

Santa Fe

Extinct

Fernandina

Very rare

Extinct7

Isabela 5 2,493

Rare to numerous

Total population

between 5,200 and 9,100. Sub-species on two south-

ern volcanoes

reduced.8

Total

13,013'

Notes:

' Townsend, 1927. These documented quantities represent only a fraction of the total amount taken.

2J. Van Denburg, 1914.

3 MacFarland, Villa, and Toro, 1974.

4 Separate races exist(ed) on each island, except Isabela which has 5.

5 Captive propagation underway to re-introduce young.

6 Eradication of introduced predators underway.

7 Last tortoise taken by a California Academy of Sciences expedition in 1905; extinction may be attributed to volcanic activity.

8 3,000 to 5,000 are found on Sierra Negra.

9 In addition, 661 tortoises have been documented as taken by scientific expeditions.

Townsend's analysis goes a step further by
estimating the total number of tortoises taken from
each island. His estimates, along with information of
the number of races found on each island and their
status in 1906 and estimated population, are
summarized in Table 2.

The last reported taking of tortoises by
whalers encountered by the author was 96 animals
taken in 1876 by the Abraham Barker of New
Bedford, but the practice surely continued into the
1900s. Townsend writes that Captain Smith, Master
of the New Bedford Bank Northern Light, while on a
voyage which passed in the vicinity of the Galapagos
in 1875 reported buying, from a small Ecuadorian
vessel sealing at the islands, 10 or 12 terrapin and a
barrel of terrapin oil, which he took home and
distributed among the housewives that he knew at
Vineyard Haven (on Martha's Vineyard,
Massachusetts).

Information on the numbers and dates of
visits by various ships frequenting the Galapagos,
along with data collected by the author confirm that,
in addition to those identified by Townsend,
approximately 1,000 visits can be documented
between 1793 and 1907. Thirty-four of these
occurred prior to 1831 and roughly 80 subsequent to
1868, so the period covered by Townsend contained
the majority of trips.

Assuming that all documented ships acted in
accordance with those identified by Townsend, at
least 70,000 tortoises were carried away. Yet, data
presented here represent only a fraction of the
American whalers that frequented the islands.
Undoubtedly, well in excess of a 100,000 were
removed and killed (the whalers were not totally
responsible, settlers also contributed). The actual

number may run into the hundreds of thousands.
The onslaught has not been totally halted. Despite
nearly 30 years of conservation efforts by the
Galapagos National Park Service and the Charles
Darwin Research Station, carcasses apparently left by
passing fishermen are still found near the coast in
remote areas of the archipelago.

Bruce C. Epler is a Marine Policy Research Assistant at the
Woods Hole Oceanographic Institution, and an international
marine economist.

Acknowledgment

The author is indebted to the staff of the Kendall Whaling
Museum for access to their valuable logs and cross-
referencing system. Julia Bryan's assistance in compiling
data is also acknowledged.

Selected References

Langdon, R. 1984. Where the whalers went: An index to the Pacific
ports and islands visited by whalers (and some ships) in the 19th
Century, pp 33-55. Canberra, Australia: Central Printery, The
Australian National University.

MacFarland, G. G., J. Villa, and B. Toro. 1974. The Galapagos Giant
Tortoises (Ceochelone elephantopus), Part 1 : Status of surviving
populations. In, Biological Conservation, Vol. 6, No. 2, April 1974.
England: Applied Science Publishers.

Shuster, G. W. 1983. The Galapagos Islands: A preliminary study of
the effects of sperm whaling on a specific whaling ground. In,
Special Issue on Historical Whaling Records: Report of the
International Whaling Commission", eds. M.F. Tillman and G. P.
Donovan, pp 81 and 82. Cambridge, England: IWC.

Starbuck, A. 1964. History of the American Whale Fishery From its
Earliest Inception to the Year 1876, Volumes I and II. New York:
Argosy-Antiquarian Ltd.

Townsend, C. H. 1927. The Galapagos Tortoises in their relation to
the whaling industry: A study of old logbooks. Zoo/og/ca, Vol. 4,
No. 1 , pp 35- 1 35. New York: The Society of the Zoological
Park.

Galapagos
Tales

For whosoever once hath fastened

His foot thereon may never it secure

But wandreth evermore uncertain and unsure.

From The Encantadas
by Salvator R. Tarnmoor

by Paul R. Ryan

Violent, tortured upheaval. This is perhaps the
single strongest image remaining from my trip to
Ecuador and the Galapagos in early January of this
year, the beginning of the rainy season. The

purpose of my visit was to prepare this issue of
Ocean us.

I left Boston's Logan Airport on New Year's
Day eve on the last flight out in the face of a major
storm moving up the East Coast. We, I was
accompanied by a colleague as far as Quito, were
bound for New York City's La Guardia airport to
make a connecting flight early the following
morning. We landed in a blinding snow storm, with
wind gusts up to 50 miles per hour. Already the
elements were setting the mood for what was to
come.

Next day, after a stop in Miami, we made
the Ecuadorian port city of Guayaquil late in the
afternoon. The plane's hydraulic system failed after
a rough landing, and we had to spend more than
three hours waiting for it to be fixed before we
could proceed to Quito, where I would spend a
couple of days on business before returning to
Guayaquil, my jumping off point for the Galapagos
Islands (Galapagos is Spanish for Tortoises). Drank
three bottles of the local Club beer in an almost
oppressive coastal heat in a Tennessee Williams-
type of airport bar with inoperable ceiling fans, and
old topographical maps of the Andes and the
Encantadas (bewitched islands) on the walls.

The 40-minute flight to Quito was
uneventful but unnerving as we flew the alley
between the snow-capped peaks of the Andes
(18,000 feet), and dropped into the colonial capital
of Quito at 9,200 feet above sea level. The lights of
the city were dazzling and "dropped into" is the
right phraseology; the landing was the hardest I've
ever experienced in a civilian aircraft, no doubt
designed to test the fragile landing-gear hydraulic
system. We were told the thinner air at that
altitude sometimes affects the depth perception of
the pilot; I felt my rear end had been only inches
from becoming a runway brake pad. I looked out
the window and thought of all the whale fossils that
were probably uplifted and buried in the
mountains outside my window:

The New York Times, Thursday,
March 12, 1987.

by Malcolm W. Browne

Scientists have found fossils of whales and other
marine animals in mountain sediments in the
Andes, indicating that the South American
mountain chain rose very rapidly from the sea.

The rare assemblage of fossils, recovered
on an expedition by the American Museum of
Natural History . . . is expected not only to
illuminate an obscure epoch of animal evolution
but also to document the rise of the Andes
mountains in the past 15 million years.

Among the fossils the scientists reported
bringing back were the bones of whales and
other marine animals found at altitudes of
more than 5,000 feet. When these animals died
from 15 million to 20 million years ago, their
carcasses settled to the ocean floor and were
embedded in submarine sediments. But since
then, the violent upthrusting of the Andean
chain has carried the sediments to the tops of

mountains. In geological terms, the time the
fossils took to rise from ocean floor to mountain
top was relatively brief.

Darwin came to the Galapagos by Beagle; Melville
in a whaleship, the Acushnet, that was his "Yale
College and . . . Harvard"; I came by TAME, an
Ecuadorian airline, along with 353 other souls,
mostly natives and a scattering of tourists. The
famous British naturalist spent 35 days in the
Galapagos, 19 of them on land, visiting 4 islands;
I'm not sure how long Melville spent while
gathering material for his short novel The
Encantadas, or Enchanted Isles; I spent 6 days,
visiting 2 islands. One of Darwin's first impressions:

They [the islands] are all formed of
volcanic rocks; a few fragments of granite,
curiously glazed and altered by the heat, can
hardly be considered as an exception. Some of
the craters are of immense size, and they rise to
a height of between 3,000 and 4,000 feet ....
Nothing could be less inviting than the first
appearance. A broken field of black basaltic
lava, thrown into the most rugged waves, and
crossed by great fissures, is everywhere covered
by stunted, sun-burnt brushwood, which shows
little signs of life. The dry and parched surface,
being heated by the noonday sun, gave to the
air a close and sultry feeling, like that from a
stove: We fancied that even the bushes smelt
unpleasant. . . .

Considering that these islands are placed
directly under the equator, the climate is far
from being excessively hot; this seems chiefly
caused by the singularly low temperature of the
surrounding water, brought here by the great
southern Polar current. Excepting during one
short season, very little rain falls, and even then
it is irregular; but the clouds generally hang low.
Hence, whilst the lower parts of the islands are
very sterile, the upper parts, at a height of a
thousand feet and upwards, possess a damp
climate and a tolerable luxuriant vegetation.
This is especially the case on the windward
sides of the islands, which first receive and
condense the moisture from the
atmosphere. . . .

This archipelago has long been
frequented, first by the buccaneers, and latterly
by whalers, but it is only within the last six years
that a small colony has been established here.
The inhabitants are between 200 and 300
hundred in number; they are nearly all people
of colour, who have been banished for political
crimes from the Republic of the Equator, of
which Quito is the capital. The settlement is
placed about 4!/2 miles inland, and at a height
of probably 1,000 feet.

Melville's first impression was somewhat different:

Take five-and-twenty heaps of cinders
dumped here and there in an outside city lot;
imagine some of them magnified into

mountains, and the vacant lot the sea; and you
will have a fit idea of the general aspect of the
Encantadas, or Enchanted Isles. A group rather
of extinct volcanoes than of isles; looking much
as the world at large might, after a penal
conflagration. . . .

The special curse, as one may call it, of
the Encantadas, that which exalts them in
desolation above Idumea and the Pole, is that
to them change never comes; neither the
change of seasons nor of sorrows. Cut by the
Equator, they know not autumn and they know
not spring; while already reduced to the lees of
fire, ruin itself can work little more upon them.
The showers refresh the deserts, but in these
isles, rain never falls. Like split Syrian gourds, left
withering in the sun, they are cracked by an
everlasting drought beneath a torrid sky. "Have
mercy upon me," the wailing spirit of the
Encantadas seems to cry, "and send Lazarus that
he may dip the tip of his finger in water and
cool my tongue, for I am tormented in this
flame. ..."

On most of the isles where vegetation is
found at all, it is more ungrateful than the
blankness of Aracama. Tangled thickets of wiry
bushes, without fruit and without a name,
springing up among deep fissures of calcined
rock, and treacherously masking them; or a
parched growth of distorted cactus trees.

In many places the coast is rock-bound,
or more properly, clinker-bound; tumbled
masses of blackish or greenish stuff like the
dross of an iron-furnace, forming dark clefts and
caves here and there, into which a ceaseless sea
pours a fury of foam; overhanging them with a
swirl of gray, haggard mist, amidst which sail
screaming flights of unearthly birds heightening
the dismal din. However calm the sea without,
there is no rest for these swells and those rocks,
they lash and are lashed, even when the outer
ocean is most at peace with itself. On the
oppressive, clouded days such as are peculiar to
this part of the watery Equator, the dark vitrified
masses, many of which raise themselves among
white whirlpools and breakers in detached and
perilous places off the shore, present a most
Plutonian sight. In no world but a fallen one
could such lands exist.

My first impressions: From several thousand
feet, as we broke out of the clouds, the islands
looked no different from those in the Caribbean or
South Pacific. But as we [the TAME jetliner] got
closer to the single runway at Baltra Airport, the
true meaning of the word desert began to take
form in my mind along with images of large cactus
trees, parched earth, and twisted, tormented
volcanic rock.

As we departed the aircraft, which was
parked a fair hike from the "airport" — a large, open
red-brick shed offering shade, Buoy and Gull
facilities, a gift shop, and a beer and soda stand—
we were herded, not unlike cattle, into lines to pay
the $40 National Park entrance fee. Natives, who
paid $40 for the roundtrip airfare to the Galapagos,

of course, passed right on through (the National
Park covers about 90 percent of the islands
territory and residents live in zones outside the
park and are therefore exempt from the tax unless
they plan to enter the park proper) to collect their
bags and ride the early, uncluttered 1950-vintage
buses to the ferry for the Island of Santa Cruz.
Tourists, naturalists, and working men like myself,
who had paid $325 for the same roundtrip ticket,
got out their handkerchiefs, passports and
Traveller's Checks or Ecuadorian sucres while
marveling at the hot, but relatively Arizona-like
climate. A woman near me, perhaps in her late 60s,
whispered to a companion: "The tour people never
mentioned an entry fee."

"I wonder if the boat will be there to meet
us," her traveling companion answered. " You hear
so many nightmare stories about tourists being
stranded in the Galapagos."

Eventually, my entry tax paid, I sought my
duffle bag — borrowed from my daughter for the
trip — with Prudhoe Bay, Alaska, stenciled on the
side. It seemed some 23 pieces of luggage were
missing — mine among them. After much desperate
arguing in pidgin-Spanish and after being told to
come back tomorrow (a 5-hour trip from where I
was staying), the head of the baggage department
was persuaded to look a final time in the plane's
baggage department, now being loaded for the
daily flight back to the mainland. Wonder of
wonders, the baggage was found.

Some 60 of us crammed into the last grime-
caked bus — probably meant to carry 40 souls
without baggage at most — to the ferry. Of course,
there are no paved roads on the island of Baltra, or
for that matter on neighboring Santa Cruz. Paved
roads, some argue, would open the door to
mainland developers, while at the same time
diminishing the nature of the Galapagos
"experience." The final descent down the cliff road
to the ferry was an exercise in pure tip-over fear.
But, as they say, "we made it!" We were next
herded onto the ferry — a launch built for 35 souls.
Our baggage loosely rode the deck roof and,
without any prospect for life preservers, we settled
back to watch the whitetip shark fins circling the
launch on its slow, 50-sucre ride across to Santa
Cruz.

At the small dock landing on Santa Cruz, we
found that the buses scheduled to take us to
Puerto Ayora — a town of some 5,000 people clear
across the island (it is estimated that 8,000 to
10,000 people live in the Galapagos) — had
returned to Puerto Ayora empty, unaware that our
baggage dilemma had delayed our arrival on Santa
Cruz. Fortunately, there was a single bus remaining
at the otherwise empty site waiting for a party of 12
that had gone out on a charter fishing boat. He
promised to take us to Puerto Ayora when his
charter returned — in about an hour's time.
However, he would not let us enter the bus until
his party was first seated for the return trip. At this
point, I befriended a young American woman who
was the daughter of a diplomat in Quito and who
was traveling to the Charles Darwin Research
Station to offer her voluntary services for the

V/A

summer months. She carried a canteen of purified
water and generously gave me a couple of thirst-
quenching sips.

Finally, we were on our way along the
straight, dirt road, pleasingly devoid of any
roadside buildings, billboards, or stands. Again, the
bus was filled to capacity, with a number of people
standing in the aisles and three passengers riding
on the roof along with most of the baggage.

At a point about half-way on the bumpy
one-and-a-half lane road — as we were climbing
I through heavy, lush green vegetation — a bag fell
off the top of the bus, breaking open and spilling
the contents of cocktail dresses, frilly underwear,
pointed shoes, and perfumes and toilet articles

•N

back down along the road for some distance. The
bus stopped, and a perplexed couple, perhaps
Americans, got out to retrieve the bag and its
contents. While retrieving their goods — the woman
wore a tight, fashionable skirt and blouse; the man
a heat-retaining business suit — the bus started up
and pulled away.

The couple, I later learned, spent 14 hours
on the road before being picked up by a bus
headed back in the opposite direction to the
airport. Once at the airport, they spent 30 hours
waiting to get space on a return flight to mainland
Ecuador. Oh, I forgot to mention, on arrival their
bag was slung off the roof into a puddle of oil.

Meanwhile, as I progressed onward, the first
Galapagos animals that I saw from the bus
windows — I saw no stores, no shops, no gas
stations — were grazing jersey cows and some
domesticated horses in farm fields. Banana tree
groves, tobacco plants, and corn plots were also
evident. About an hour out of Puerto Ayora, the
bus stopped for bladder drains and a look at two
deep volcano craters.

It was about 7 p.m. when we pulled into the
town's main square near the harbor. It was past
sunset but with still enough light to see the cactus
and generally parched terrain. The "boom town"
nature of Puerto Ayora — the largest metropolis in
the Galapagos and the hub of the tourist trade-
was readily apparent. The streets were unpaved
and dusty, and the one-story building facades
chipped and cracked, the once bright wall colors
bleached dull by a relentless sun. New
construction starts here and there dotted the
streets — their tell-tale piles of brick and bags of
cement stacked on the sidewalks. Children smiled
engagingly from entrance ways. I noted a pizza
parlor.

It was a 30-minute hike with bags (there
were no taxis in the town) to my first-class hotel—
"the Galapagos," owned by an American who
happened, at this moment, tp be vacationing by
motorcycle in the jungles of Thailand. I was
impressed by the poisonous fruit tree in the
courtyard. As I was shown my cottage, I was
assured I would see large black spiders on the wall
occasionally, but not to worry, they were harmless
and preyed on the large (2-inch) flying cockroaches
that sometimes put in an appearance. The gentle
pounding of the sea on lava rocks outside my
window was a reassuring sound. The town
generator, I was told, would shut off at midnight,
and anyone wanting to read or walk would have to
light candles or a flashlight.

The bar was generally unattended, guests
being introduced to the honor system. Beer, cokes,
7-up, and Gitig (a local mineral water) were
available, along with a limited rum, gin, and
whiskey supply. One signed a chit at the bar after
mixing one's own drink. While I was there, it was
seldom patronized after 9 p.m.

There were six people staying at the
Galapagos Hotel when I arrived. Four of them were

frequent visitors to Woods Hole, including one
summer resident of Penzance Point and two
subscribers to Oceanus. It is indeed a small world.
They were all naturalists of one ilk or another, four
of them already hardened by the rigors of
photographic safaris in East Africa.

In the morning, a cow bell, gently rung,
announced breakfast at 7 a.m., which was served
until 7:30 a.m. on six large picnic tables in the main
lobby. Coffee, fruit, pancakes, sometimes eggs or
bacon, and juice was the main fare. I was greeted
on the paved cement path to breakfast by two sun-
bathing marine iguanas. Yes, I must be in the
Galapagos after all, I remember thinking, marveling
at these magnificent beasts who stirred memories
of the dinosaur periods. They seemed not to mind
my presence in the least.

* * *

Later that morning, I stumbled on a group of
marine iguanas on the black lava rocks behind the
hotel. This was an area that served as a back lawn
with short, stubby vegetation among the rocks
down to the sea.

It was breeding season and two large males
were butting heads in a contest for territorial rights.
I raced back to my cottage and grabbed my
camera, borrowed from the Woods Hole
Oceanographic Institution for the trip. The shot
counter was on 10 and I felt lucky that I would not
have to load film.

I got back in time to witness the end of the
head-butting contest, getting some wonderful
shots. I was impressed by the speed of the two
iguanas when head butting. They would circle each
other and then, in a lightning-fast move, bang
heads. Finally, one of the iguanas, looking a little
dazed, backed off. The other large iguana shortly
chose a smaller female iguana and began the
mating process, which lasted a good 20 minutes.

Three baby iguanas, perhaps six inches long,
watched from a perch on a rock above the burrow
where the pair was mating. I thought I had some
world-class shots of this type of behavior. Later,
upon opening my camera, I discovered that there
was, indeed, no film in it.

The Galapagos are home for a number of European
exiles. The Angermeyers fall into this category. Carl
and his two brothers left Germany as youths in
1937 to take up residence. They hunted wild goats,
caught rock lobsters and groupers, and farmed.
Carl, a boat captain and artist among his many
talents, has given shelter to a large number of
marine iguanas in his home. Angermeyer's pets
have developed tastes for more than just seaweed
and algae, they also enjoy boiled rice, pancake
batter, and raw fish from one's fingers.

* * *

Came across this item in my nighttime reading in
the Galapagos. It seems a group of Americans led
by a "casual adventurer and scientific fiction
reader" founded a colony in 1959 on San Cristobal
Island for the purpose of exploiting the island's
resources.

R. E. L. Paris and others, writing in the Pacific
Sociological Review, tell us that the leader's ideal of
society was one based on "clear thinking and
scientific principles."

The Ga/apagos Islands were perceived by
the leader to be rich in marketable resources. At
various times he made mention of the
possibilities of profit from coffee-growing, cattle-
raising, lobster-fishing, seaweed-gathering,
tourist-entertaining, and scientific research,
especially biological studies. The plan of
organization known as "Filiate Science
Antrorse" (meaning together with science we
move forward) did not take into serious
consideration the Ecuadorian residents of the
islands. The final document of organization of
F.S.A., however, lured more than 100 persons to
the islands, representing 36 family units, each of
whom turned over $2,500 of their savings,
committing their future lives to this scheme.

Most of the recruits were drawn from
applicants living in the State of Washington.
They were chiefly young persons with moderate
incomes, including aircraft workers, farmers,
truck drivers, firemen, salesmen, a janitor, a
plumber, and some school teachers. One
feature they all seemed to have in common was
a dissatisfaction with their present condition of
life, a mixture of idealism, and a yearning for a
new and more exciting direction in a so-far
adventure/ess career.

A series of disappointments, including
lack of seaworthiness of their ship, the Alert,
lack of fishing skills among its members,
depletion of the local lobster resource,
irreparability of the refrigeration plant at Puerto
Baquerizo Moreno, unavailability of their

hoped-for coffee-plantation in the highlands of
San Cristobal, political troubles in Ecuador
resulting from this "Yankee Invasion," and
debilitating diseases such as dysentery and
hepatitis, resulted in a total collapse of the
venture. By January, 1961, almost 14 months
after the first group of colonists had reached San
Cristobal, all but one of the original colonizers
had left the Galapagos. Thus, in little more than
a year, 106 persons had come and gone from
their Utopian island, spent an estimated
$165,000, experienced personal bankruptcy,
and become generally disillusioned.

— from Contributions to Science from the
Galapagos by Robert I. Bowman, Key
Environments, 1984.

* * *

One morning I visited Tortuga Bay in a small
launch that left from the hotel dock. A large brown
pelican perched calmly on a piling observed our
departure. There were nine in the party, including
an American naturalist guide who was born in the
islands and who operated a cattle and vegetable
farm when not working as a tour guide. It was
about an hour and a half down the coast to the
bay. As we left, we got a good view of the
glistening visiting yachts and local touring
schooners moored in the harbor outside Puerto
Ayora.

Once outside the reefs, the sea was choppy.
Saw two giant turtles at a distance. On entering
Tortuga Bay, I saw a seven-foot whitetip shark
break water in a thrashing motion. He was in
shallow water, probably feeding. Colleagues had
advised me before I left that 1) sharks were
plentiful in Galapagos waters; 2) none had ever
killed a tourist, although native commercial divers
had been known to disappear from time to time; 3)
it was not uncommon for bull and other sharks to
charge and nudge divers on occasion; and 4) the
watchword was to be wary. I kept near the launch
while doing some snorkling in Tortuga Bay. The
water was turbid and one could only see about
three feet. Did manage to get within two feet of a
blue-footed boobie, however, who seemed very
interested in observing the behavior of our group.
Did not see any turtles in Tortuga Bay but did see a
couple of magnificent flamingos, a lovely white
fine-sand beach, and two cans filled high with
human visitors' trash.

* * *

I was disappointed that the owner of the Galapagos
Hotel was on vacation. He is an avid chess player,
as I am whenever I get the chance, which is not
often. The hotel owner has a human-size board
where chess pieces are moved by long shuffle
board-like sticks from two king-size lifeguard chairs
at either end of the board.

My six-day visit came to an end looking for
XL teeshirts to take home. The shops were small
but numerous along the road to the center of town
and full of black coral jewelry items, despite the
fact that it is reportedly illegal to harvest black
coral. One shop owner explained: "It may be illegal
to harvest it, but it is not illegal to make jewelry
from it."

While the plane ride back to Woods Hole
was uneventful, I should mention that a military C-
145 crammed with Ecuadorian troops took off from
Quito for Guayaquil shortly before my Eastern
flight to Panama. Before we would arrive in
Panama, Ecuadorian Air Force commandos would
seize President Leon Febres Cordero, a 54-year-old
millionaire businessman, as a hostage in exchange
for the release of an Air Force general being held in
custody for two uprisings against the Defense
Minister despite a Congressional amnesty. Just
another upheaval in a land of upheavals.

* * *

The Galapagos Islands have the image among those
who read the ads and articles in slick magazines
around the world of being a Pacific paradise, a
place to visit on a luxurious cruise boat at some
point in life when one is embarked on seeing the
remote wonders of the world. On such a cruise
one can enjoy all the amenities of life while
vicariously bearing witness to the realities.

The Galapagos Islands are not a paradise in
the commercial advertising sense of the word,
although they very well may be "treasure islands"
to scientific researchers. This raises the question of
tourism, which is the single greatest management
issue for the newly created Galapagos Marine
Resources Reserve (see Broadus article page 9)—
the raison d'etre for this issue. At present, the
islands entertain more than 25,000 tourists a year
with a population of from 8,000 to 10,000 residents
scattered over the four inhabited islands in the
archipelago. A battle is shaping up between
conservationists who are defending the unique and
pristine qualities of "Darwin's laboratories" on the
one hand, and the developers who want to open
the islands to high-rise hotels, 150,000 tourists a
year, and the Pacific paradise image on the other.
This, of course, would help to make them and
Ecuador richer.

But there is another disturbing factor in the
equation. Some argue that the residents of the
islands, many of whom live on the brink of survival
and who could use a large measure of fitness,
would not benefit from increased tourism — that
most of any newly created jobs would go to
persons with hotel and tour experience on
mainland Ecuador. It is not, for this writer, a
convincing argument. Whatever the decision on
development of the islands, attention must be paid
to the plight of the people, both those who live in
the archipelago and those in mainland Ecuador on
an equal basis.

Probably the best idea I heard during my
visit was a suggestion that mainland Ecuador has
many undeveloped areas of tourist interest
comparable to those of the Galapagos. Why not
develop these attractions, thereby taking the
pressure off the Galapagos to accommodate more
tourists and leaving the islands as a "paradise" lost
to science, nature, volcanism, and time.

Paul R. Ryan is Editor of Oceanus, published by the Woods
Hole Oceanographic Institution.

Acknowledgment

Sketches by Sig Purwin, Woods Hole, MA.

Suhtidal Galapagos by James Cribb. 1986. Camden House
Publishing Ldt, Ontario, Canada. 153 pp. $29.95.

Three major ocean currents — the cool fertile Humboldt,
the warm tropical El Nino and the Cromwell, a subsurface
current that carries nutrient-rich waters — converge in the
Galapagos to produce an underwater world which rivals its
more publicized terrestrial counterpart in uniqueness and
diversity. James Cribb sets out to explore this fascinating,
little known marine environment, and to bring the reader
along on his adventure.

Introductory chapters are dedicated to identifying
some of the logistics involved in coordinating such an
undertaking, and a brief but clear description of the
complex oceanographic and climatic factors that are
responsible for the coexistence of subantarctic and tropical
life forms in waters surrounding the archipelago. The
remaining chapters present accounts of voyages through
four marine provinces. The journey begins by visiting the
central islands with their broad representation of marine
creatures, continues on to the cool waters and plunging
volcanic cliffs of the western islands, and then to the shark-
infested waters surrounding the southern islands. The last
area visited is the northern islands, where tropical species
including coral abound.

The well-written and often exciting narrative is
greatly enhanced by 117 color photographs. The
extraordinary collection of marine creatures photographed
are a testimony to the beauty and brilliant colors to be
found in this extraordinary world and leave the reader with
an appreciation of both the Galapagos marine and
terrestrial environments. Publication of the book is timely as
the interior waters of the archipelago and a 15-mile buffer
zone were decreed a Marine Resources Reserve in May,
1986.

The publication could be improved by better
coordination of text and photographs as the reader is often
caught up in a description but unable to find a
corresponding photograph.

Anyone interested in learning more about the
Galapagos marine environment without getting into
scientific nomenclature will find Subtidal Galapagos
appealing. The book is a must for those contemplating a
diving trip to the islands and a valuable addition to any
browsing table or collection of nature books.

Bruce Epler,
Research Assistant,

Marine Policy and Ocean Management Center,
Woods Hole Oceanographic Institution

Deep-Sea Challenge: The ]ohn Murray /Mabahiss
Expedition to the Indian Ocean, 1933-34, A. L. Rice, ed.
1986. UNESCO Press, Paris. Distributed in the U.S. by
Bernan Associates-UNIPUB, Lanham, MD. 336 pp. $30.00.

While perusing the book stacks in the library of the Marine
Biological Laboratory some 40 years ago, trying to educate
myself in oceanography, I found that the decade 1915-
1935 seemed to mark the puberty of physical ocean-
ography. There, I encountered expedition reports of
research ships such as the Meteor, Discovery, Dana, and
Snellius, and became especially interested in the
1 1 volumes of the John Murray Expedition, 1933-1935 and
the story behind it.

Sir John Murray, the scientific editor of the HMS
Challenger Reports, had discovered commercially valuable
phosphate in rock samples brought to him from Christmas
Island in the Indian Ocean, and with the help of a local
plantation owner, George Clunies Ross, "King of the
Cocos," had exploited the deposits very profitably. He
made enough money to establish a fund for research after
his death in 1914. The fund grew with time, and it was
decided to sponsor an oceanographic expedition to the
northwest Indian Ocean. The Egyptian government loaned
the expedition their newly acquired steam vessel Mabahiss;
the fund paid for salaries of officers, crew, and scientists, all
running costs such as food and coal, shipping, travel
expenses, and scientific equipment, for the nine months of
the expedition, and for publication of the results. The total
budget was £20,000. (It would probably cost 100 times that
today: a factor of 20 through inflation, and a factor of five
because it always costs more for the government to sponsor
anything.)

During a lecture stint at Yale in the 1950s, I came to
know E. F. (Bill) Thompson, who had been the physical
oceanographer on the Murray expedition. He also had
been responsible for the cruises of the Royal Society's
vessel, Culver, off Bermuda that followed. He had access to
the working papers for both these field studies, but
unfortunately never pursued their publication. I considered
this a loss to the scientific community, as the results of the
Murray expedition had always seemed to me to be only
partly written up.

However, the leader of the John Murray Expedition,
Lt. Col. R. B. Seymour Sewell evidently left a typescript
version of a journal that he kept during the cruise. This
forms the core of Deep-Sea Challenge, and a very good

100

"*• AA/-
*****

Unesco

book it is. Included are: (a) a good account of the origin of
the expedition by A. L. Rice, (b) biographical notes on the
chief participants, by Rice with the help of S. A. Morcos,
(c) the history of the Mabahiss herself, and (d) a scientific
appreciation of the results of the expedition by Rice and Sir
George Deacon. There also are some rather wonderful
photographs.

The reader will avoid a shock on page 74 if he is
prepared to substitute Figure 35 on page 258 for Figure 12.
On the whole, the book is nicely produced, informative,
and fascinating reading for those who take an interest in
oceanic research of past generations.

Henry Stommel,

Senior Scientist,

Woods Hole Oceanographic Institution

EDITOR'S NOTE: The late Sir George Deacon was the
subject of a profile in Oceanus Vol. 28, No. 1; Henry
Stommel was profiled in Oceanus Vol. 27, No. 1.

The Ocean of Truth: A Personal History of Global
Tectonics by H. W. Menard. 1986. Princeton University
Press, Princeton, NJ. 353 pp. $29.50.

To a beginning student of marine geology, the theory of
plate tectonics must appear to be such a smoothly
operating mechanism that he or she has little doubt of its
ability to coordinate many different aspects of the rocks
and sediments of the Earth. In fact, the student may find it
difficult to conceive of the many fits and starts, changes of
concepts, and searches for new tests that occurred during
the main formative years, 1961-1966.

Menard was one of a score of principal investigators
whose efforts led to general acceptance of plate tectonics.

These investigators included some who worked nearly
alone and others who were aided by many assistants. Prior
to 1961, knowledge of the ocean floor was mainly
descriptive and had been gained through use of what now
would be viewed as quaint methods and equipment. The
need for broad generalizations about the origin of oceans
and continents was recognized, and preliminary efforts
were made, but with little success, owing to insufficient and
inadequately coordinated data. The real push began with
short articles by R. S. Dietz and H. H. Hess that forecast the
modern concept of sea-floor spreading, but could not
prove it. These articles essentially by-passed large gaps in
knowledge to link existing large pieces of knowledge in a
reasonable but new way.

Menard's book is a summary of subsequent efforts
to learn whether and how sea-floor spreading occurs. He
notes many of the complications that resulted from
multiple-simultaneous discoveries, rejection of manuscripts
by journal editors as too controversial or too different,
classification of some data by the U.S. Navy, and strong
positions taken by some investigators who had access to
data of only one sort or from only one ocean, and who
radically changed their views when new data became
available. The most influential investigators were from
Scripps Institution of Oceanography, Lamont Geological
Observatory, Princeton University in the United States, and
from Cambridge University in England. Investigators at
other organizations had lesser roles, and those in most
other countries made essentially no contribution during the
formative years of sea-floor spreading.

Data of many kinds eventually were incorporated in
the study, but of course they usually were not available at
the times of greatest need — thus accounting for many of
the delays and uncertainties during the investigations. Most
of the background data were available from previous long-
term shipboard studies supported mainly by the U.S. Office
of Naval Research and the National Science Foundation
through grants and contracts to investigators at
oceanographic organizations. Funds for new critical data,
and for syntheses with previous data were provided by
these same funding agencies. The kinds of information
sought ran the gamut of geology and geophysics:
physiography (continental rises, trenches, mid-ocean ridges,
transform faults, oceanic islands), distribution and depths of
earthquake epicenters, kinds of rock on ocean floors and
their ages, magnetic reversal anomalies imprinted on
cooling igneous rocks, measurements of gravity, inferences
about composition and movements of underlying mantle
rocks, thicknesses, compositions, and ages of overlying
marine sediments, and supplementary paleomagnetic
orientations of mainly continental rocks. Special tools for
measurements of seismic profiles, magnetics, gravity, and
especially deep-sea drilling had to be invented or be much
improved. Increased precision in dating of cores by
isotopes and paleontology also occurred.

By the end of 1966, the concept of sea-floor
spreading had been well established at least in the minds of
those who had done most of the work. The much
improved methods continued to be used to investigate
details and new secondary questions, and these efforts
have spread far more widely than the efforts during the few
years of the greatest progress in understanding. This is
typical of revolutions in science and it can last for a century
or more, as illustrated by the similar revolution in biology
led by Charles Darwin more than a century earlier.

Marine geology is much indebted to Bill Menard for
his recording of the uncertain and indirect evolution of the
broadest generalization of geology, a field that is little more
than a century old. During the score of years since 1 966
some of the most active investigators have died, and more

101

will go during the next decade, but Menard's summary has
captured the essence of the revolution. His summary of
conversations, meetings, letters, publications, and
personalities is an effective history of a scientific revolution
available for consultation by those who might otherwise

believe that a scientific revolution is more smooth,
reasonable, and orderly than a political one.

K. O. Emery,

Scientist Emeritus,

Woods Hole Oceanographic Institution

Books Received

Aquaculture

Shellfish and Seaweed Harvests of
Puget Sound by Daniel P. Cheney
and Thomas E. Mumford, Jr. 1987.
University of Washington Press,
Seattle, WA. 164 pp. + xv. $8.95.

Atmospheric Science

Atmospheres and Ionospheres of the
Outer Planets and Their Satellites by
Sushil K. Atreya. 1986. Physics and
Chemistry in Space 15. Springer-
Verlag, New York, N.Y. 224 pp. +
xiii. $69.50.

The Ceaseless Wind: An Introduction
to the Theory of Atmospheric Motion
by John A. Dutton. 1986. Dover
Publications, Inc., New York, N.Y.
617pp. + xix. $16.95.

The Global Climate, John T.
Houghton, ed. 1984. Cambridge
University Press, New York, N.Y. 233
pp. + v. $19.95.

Biology

The Biological Chemistry of Marine
Copepods, E. D. S. Corner and S. C.
M. O'Hara, eds. 1986. Oxford
University Press, New York, NY. 349
pp. + x. $73.00.

Caribbean Reef Invertebrates by
Nancy Sefton and Steven K. Webster.
1986. Sea Challengers, Monterey,
CA. 112pp. $19.95

Contemporary Studies on Fish
Feeding, Charle A. Simenstad and
Gregor M. Cailliet, eds. 1986. Dr. W.
Junk Publishers, The Netherlands.
334pp. $122.00.

Crabs of Cape Cod by Stephan
Berrick. 1986. The Cape Cod
Museum of Natural History,
Brewster, MA. 77 pp. $6.95.

Fish Processing in Africa:
Proceedings of the FAO Expert
Consultation on Fish Technology in
Africa. 1986. FAO Fisheries Report
No. 329. Distributed by Bernan-
UNIPUB, Lanham, MD. for the Food
and Agriculture Organization, Rome.
474 pp. + vii. $29.00.

The Fish Resources of the Northwest
Pacific by S. Chikuni. 1985.
Distributed by Bernan-UNIPUB,
Lanham, MD. for the Food and
Agriculture Organization of the
United Nations, Rome. 190 pp. + xiii.
$11.00.

Giants of Land, Sea, and Air: Past
and Present by David Peters. 1986.
Sierra Club Books, San Francisco,
CA. 73pp. $12.95.

Marine Mammals, Delphine Haley,
ed. 1986. Second edition. Pacific
Search Press, Seattle, WA. 295 pp.
$22.95.

Light and Photosynthesis in Aquatic
Ecosystems by John T. O. Kirk. 1986.
Cambridge University Press, New
York, NY. 401 pp. + xii. $24.95.

Seasonality of Freshwater
Phytoplankton, M. Munawar and J. F.
Tailing, eds. 1986. Developments in
Hydrobiology 33. Dr. W. Junk
Publishers, The Netherlands. 236 pp.
+ viii. $95.50.

1984 Yearbook of Fishery Statistics:
Catches and Landings. 1986.
Distributed in the U.S. by Bernan-
UNIPUB, Lanham, MD. for the Food
and Agriculture Organization of the
United Nations, Rome. 452 pp. + viii.
$35.50.

7984 Yearbook of Fishery Statistics:
Fishery Commodities. 1986.
Distributed in the U.S. by Bernan-
UNIPUB, Lanham, MD. for the Food
and Agriculture Organization of the
United Nations, Rome. 310 pp. + vii.
$27.00.

Earth Sciences

Advances in Soil Science: Vol. 6, B.
A. Stewart, ed. 1987. Springer-
Verlag, New York, NY. 222 pp. + viii.
$64.70.

Global Bio-Events, Otto H. Walliser,
ed. 1986. Springer-Verlag, New
York, N.Y. 442 pp. + vii. $38.50.

The Indian Ocean: Exploitable
Mineral and Petroleum Resources by
G. S. Roonwal. 1986. Springer-
Verlag, New York, N.Y. 198 pp. + xv.
$61.00.

Oceanology of the Antarctic
Continental Shelf, Stanley S. Jacobs,
ed. 1985. Antarctic Research Series
43. American Geophysical Union,
Washington, D.C. 312 pp. + ix.
$39.00.

Offshore Seismic Exploration by
Rajni K. Verma. 1986. Gulf
Publishing Co., Houston, TX. 591 pp.
+ xiv. $75.00.

Sediments and Water Interactions,
Peter G. Sly, ed. 1986. Springer-
Verlag, New York, N.Y. 521 pp. +
xxi. $105.00.

The Superdeep Well of the Kola
Peninsula, Ye. A. Kozlovsky, ed.
1987. Springer-Verlag, New York,
NY. 558pp. + xi. $118.00.

Ecology/Environment

The Background of Ecology: Concept
and Theory by Robert P. Mclntosh.
1986. Cambridge University Press,
New York, NY. 383 pp. + xiii.
$16.95.

Biological Processes and Wastes in
the Ocean, Judith M. Capuzzo and
Dana R. Kester, editors. 1987.
Oceanic Processes in Marine
Pollution 1. Robert F. Krieger
Publishing Co., Malabar, FL. 265 pp.
+ xiii. $43.50.

Caribbean Coastal Marine
Productivity: Results of a Planning
Workshop at Discovery Bay Marine
Laboratory, University of the West
Indies, Jamaica. 1986. Unesco, Paris.
59 pp. free.

Coastal Off-Shore Ecosystems
Relationships. Final Report of SCOR/
lABO/Unesco Working Group 65.
1986. Unesco, Paris. 39 pp. + vi.
free.

102

The Disposal of Long-Lived and
Highly Radioactive Wastes, A. S.
Laughton, L. E. ). Roberts, Denys
Wilkinson, and D. A. Gray, eds.
1986. The Royal Society, London.
189 pp. + v. £ 33.00.

The Ecology of River Systems, B. R.
Davies and K. F. Walker, eds. 1986.
Dr. W. Junk Publishers, The
Netherlands. 793 pp. + xviii.
$148.00.

Evolution

The Correspondence of Charles
Darwin, Frederick Burkhardt and
Sydney Smith, eds. 1986. Volume 2.
Cambridge University Press, New
York, NY. 603 pp. + xxxiii. $37.50.

The Darwinian Heritage, David
Kohn, ed. 1985. Princeton University
Press, Princeton, NJ. 1 138 pp. + xii.
$95.00.

Patterns and Processes in the History
of Life, D. M. Raup and D. Jablonski,
eds. 1986. Life Sciences Research
Report 36. Springer- Verlag, New
York, N.Y. 447 pp. + xi. $88.00.

Field Guides

California Marine Food and Game

GALAPAGOS

You, 9 other
adventurers and

our licensed

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INCH 6FLOHTS

I606on Juanita. Tiburon, CA 94920
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Fishes by John E. Fitch and Robert |.
Lavenberg. 1971. University of
California Press, Berkeley. 179 pp.
$5.95.

Dive to the Coral Reefs by Elizabeth
Tayntor, Paul Erickson, and Les
Kaufman. 1986. Crown Publishers,
Inc., New York, N.Y. 36 pp. $12.95.

Northwest Shore Dives by Steve
Fischnaller. 1986. Bio-Marine Press,
Edmonds, WA. 240 pp. $12.95.

Reef Fishes of the Sea of Cortez by
Donald A. Thompson, Lloyd T.
Findley, and Alex N. Kerstitch. 1987.
The University of Arizona Press,
Tucson, AZ. 302 pp. + xviii. $19.95.

Tidepool and Nearshore Fishes of
California by John E. Fitch and Robert
J. Lavenberg. 1975. University of
California Press, Berkeley. 156 pp.
$3.95.

Treasures of the Tropic Seas by Rene
Catala. 1986. Facts On File, Inc., 460
Park Avenue South, New York, N.Y.
334 pp. $50.00.

General Reading

Development of Marine Sciences in
Arab Universities: Meeting of Experts
held at the Marine Science Station in

Aqaba, Jordan 1-5 December 1985.
1986. Unesco Reports in Marine
Science 39. Unesco, Paris. 58 pp.
free.

Essentials of Ocean Science by Keith
Stowe. 1987. John Wiley and Sons,
Inc., Somerset, N.J. 353 pp. + xi.
$33.55.

Fjords: Processes and Products by
James P. M. Syvitski, David C.
Burrell, and Jens M. Skei. 1987.
Springer-Verlag, New York, N.Y. 379
pp. + x. $85.00.

Oceanography: A View of the Earth
by M. Grant Gross. 1987. Fourth
Edition. Prentice-Hall, Inc.,
Englewood Cliffs, NJ. 406 pp. + ix.
$35.33.

The Scientist at the Seashore by
James S. Trefil. 1987. Macmillan
Publishing Co., New York, NY. 224
pp. $8.95.

Information and
Directories

International Directory of Marine
Science Libraries and Information
Centers. Compiled by Carolyn P.
Winn. 1987. Woods Hole
Oceanographic Institution and
International Association of Marine

For moored, fixed position, or profiling
measurement of temperature and
salinity at depths to 6800 meters:

SEA-BIRD'S new SEACAT and SEACAT PROFILER
offer proven Sea-Bird conductivity and
temperature sensors in a self-contained solid
state logging package.

These compact instruments provide very
high resolution and accuracy flexible
acquisition routines, precision time-
bases, and convenient electrical read-
out. Pressure measurement available
in SEACAT, standard in SEACAT
PROFILER.

The best sensors are even better.

Manufacturers of cable telemetering and internal recording LTD systems, modular sensors and data loggers for
conductivity, temperature, dissolved oxygen. pH. and other environmental variables

SE*E Sea-Bird Electronics, Inc.

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Telephone: (206) 462-8212. Telex: 292915 SBEI UR.

103

Science Libraries and Information
Centers, Woods Hole, MA. $15.00.

Marine Policy

The Beaches are Moving: The
Drowning of America's Shoreline by
Wallace Kaufman and Orrin H.
Pilkey, Jr. 1984. Living with the Shore
1. Duke University Press, Durham,
NC. 336 pp. + viii. $9.75.

The Law of the Sea: Master File
Containing References to Official
Documents of the Third United
Nations Conference on the Law of
the Sea, by the Office of the Special
Representative of the Secretary-
General for the Law of the Sea. 1985.
United Nations, New York, N.Y.
Distributed by Bernan Associates-
UNIPUB, Lanham, MD. 176 pp.
$19.50.

The Law of the Sea, Pollution by
Dumping: Legislative History of
Articles 1, Paragraph 1(5), 210 and
21 6 of the United Nations
Convention on the Law of the Sea,
by the Office of the Special
Representative of the Secretary-
General for the Law of the Sea, 1985.
United Nations, New York, N.Y.
Distributed by Bernan Associates-
UNIPUB, Lanham, MD. 77 pp.
$11.50.

The Law of the Sea Treaty: One
Observer's Assessment of the
Conference, the Treaty and Beyond
by Thomas A. Clingan, Jr. 1986.
Washington Sea Grant Program.
Distributed by Washington Sea Grant
Communications, Seattle, WA. 19 pp.
$3.00.

Living with the Coast of the Puget
Sound and Georgia Strait by Thomas
A. Terich. 1987. Living with the
Shore. Duke University Press,
Durham, NC. 165 pp. + xv. $12.95.

Methodologies for Assessing the
Impact of Deep Sea-Bed Minerals on
the World Economy by the
Department of International
Economic and Social Affairs. 1986.
United Nations, New York, NY. 153
pp. + ix. $16.50.

Natural Resources Economics and
Policy Applications: Essays in Honor
of James A. Crutchfield, Edward
Miles, Robert Pealy, and Robert
Stokes, eds. 1986. University of
Washington Press, Seattle, WA. 456
pp. + xiii. $30.00.

Ocean Forum: An Interpretative
History of the International North
Pacific Fisheries Commission by Roy
I. Jackson and William F. Royce.

1986. Fishing News Books Ltd.,
Surrey, England. Distributed in the
U.S. by Bernan-UNIPUB, Lanham,
MD. 240pp. $31.50.

Ordering the Oceans: The Making of
the Law of the Sea by Clyde Sanger.

1987. University of Toronto Press,
Toronto, Ontario. 225 pp. + xii.
$14.95.

Physical Sciences

Acoustic Waves: Devices, Imaging,
and Analog Signal Processing by
Gordon S. Kino. 1987. Prentice Hall,
Inc., Englewood Cliffs, NJ. 601 pp. +
xxi. $64.00.

The Application of Digital Remote
Sensing Techniques in Coral Reef,
Oceanographic and Estuarine
Studies: Report on a regional
Unesco/COMAR/GBRMPA
Workshop in Townsville, Australia.
1986. Unesco, Paris. 59 pp. free.

Environmental Hydraulics: Stratified
Flows, Malcolm J. Bowman, Richard
T. Barber, Christopher N. K. Mooers,
and John A. Raven, eds. 1986.
Springer- Verlag, New York, N.Y. 278
pp. + xv. $31.90.

Geophysical Fluid Dynamics by
Joseph Pedlosky. 1987. Second
edition. Springer- Verlag, New York,
N.Y. 710 pp. + xiv. $49.00.

General Circulation of the Ocean,
Henry D. I. Abarbanel and W. R.
Young, eds. 1987. Springer- Verlag,
New York, N.Y. 291 pp. + xii.
$69.00.

Hydrodynamics of Ocean Wave-
Energy Utilization, D. V. Evans and
A. F. de O. Falcao, eds. 1986.
Springer- Verlag, New York, N.Y. 452
pp. + xvi. $49.00.

The Physical Nature and Structure of
Oceanic Fronts by K. N. Fedorov.
1986. Lecture Notes on Coastal and
Estuarine Studies 19. Springer-
Verlag, New York, N.Y. 333 pp. +
viii. $69.00.

Physics of Shallow Estuaries and
Bays, J. van de Kreeke, ed. 1986.
Springer-Verlag, New York, N.Y. 280
pp. + vii. $24.50.

Thermal Modeling in Sedimentary
Basins, Jean Burrus, ed. 1986. IFP
Exploration Research Conference 44.
Gulf Publishing Company, Houston,
TX. 600 pp. -I- xix. $89.00.

Topics in Geophysical Fluid
Dynamics: Atmospheric Dynamics,
Dynamo Theory, and Climate
Dynamics, by M. Ghil and S.
Childress. 1987. Applied
Mathematical Sciences 60. Springer-
Verlag, New York, NY. 485 pp. + xv.
$39.00.

Science Communication

The Visual Display of Quantitative
Information by Edward R. Tufte.
1983. Graphics Press, Cheshire, CT.
197 pp. $32.00.

Ships and Sailing

Ancient Boats in N.W. Europe by
Sean Me Grail. Longman, New York,
NY. 1987. 321 pp. + xx. $79.95.

Arctic Whalers, Icy Seas: Narratives
of the Davis Strait Whale Fishery by
W. Gillies Ross. 1985. Irwin
Publishers, Toronto, Canada. 263 pp.
+ xvi. $25.95.

The Battleship Warspite by Ross
Watton. 1986. Anatomy of the Ship
9. Naval Institute Press, Annapolis,
MD. 120pp. $21.95.

Captains of the Old Steam Navy,
James C. Bradford, ed. 1986. The
Naval Institute Press, Annapolis, MD.
356 pp. + xvi. $24.95.

A Cruising Guide to the New
England Coast by Roger F. Duncan
and John P. Ware. 1987. Ninth
Edition. Dodd, Mead and Company,
New York, NY. 732 pp. + xviii.
$12.95.

Fleet Tactics: Theory and Practice by
Capt. Wayne P. Hughes, Jr. 1986.
The Naval Institute Press, Annapolis,
MD. 316 pp. + xvi. $21.95.

Fletcher-Class Destroyers by Alan
Raven. 1986. The Naval Institute
Press, Annapolis, MD. 158 pp.
$21.95.

Guide to the Soviet Navy by Norman
Polmar. 1986. Fourth Edition. The
Naval Institute Press, Annapolis, MD.
536 pp. + xii. $38.95.

The Last Navigator by Stephen D.
Thomas. 1987. Henry Holt and Co.,
New York, NY. 308 pp. $22.95.

Nautical Quarterly: No. 37, Spring
1987. Nautical Quarterly Co., Essex,
CT. 124pp. $16.00.

The Seventy-Four Gun Ship: Vol. 1,
Hull Construction by Jean Boudriot.
Naval Institute Press, Annapolis, MD.
166pp. $58.95.

104

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Department of International
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Natural Resources Economics and
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Miles, Robert Pealy, and Robert
Stokes, eds. 1986. University of
Washington Press, Seattle, WA. 456
pp. + xiii. $30.00.

Ocean Forum: An Interpretative
History of the International North
Pacific Fisheries Commission by Roy
I. Jackson and William F. Royce.

Oceanic Fronts by K. N. Fedorov.
1986. Lecture Notes on Coastal and
Estuarine Studies 19. Springer-
Verlag, New York, N.Y. 333 pp. +
viii. $69.00.

Physics of Shallow Estuaries and
Bays, J. van de Kreeke, ed. 1986.
Springer-Verlag, New York, N.Y. 280
pp. + vii. $24.50.

Thermal Modeling in Sedimentary
Basins, Jean Burrus, ed. 1986. IFF
Exploration Research Conference 44.
Gulf Publishing Company, Houston,
TX. 600 pp. + xix. $89.00.

Guide to the Soviet Navy by Norman
Polmar. 1986. Fourth Edition. The
Naval Institute Press, Annapolis, MD.
536 pp. + xii. $38.95.

The Last Navigator by Stephen D.
Thomas. 1987. Henry Holt and Co.,
New York, NY. 308 pp. $22.95.

Nautical Quarterly: No. 37, Spring
1987. Nautical Quarterly Co., Essex,
CT. 124pp. $16.00.

The Seventy-Four Gun Ship: Vol. 1,
Hull Construction by Jean Boudriot.
Naval Institute Press, Annapolis, MD.
166pp. $58.95.

104

LIBRARY

Oceanus

Japan

and the Sea

Vol. 30:1, Spring 1987— The
first comprehensive view of
Japanese ocean science
written primarily by Japa-
nese authors. Describes how
tradition and innovation
combine to continue forging
a strong link between Japan
and the sea. Includes fishing,
submersibles, SWATH ves-
sel, recovery of uranium,
ocean space, and much,
much more.

Changing Climate
and the Oceans

Vol. 29:4, Winter 1986/87-
Forecasts of near-term cli-
mate change have chal-
lenged scientists to under-
stand complex interactions
between the atmosphere,
the ocean, and the Earth.
The wobbling Earth, chang-
ing sunlight, carbon dioxide,
polar ice sheets, and defo-
restation— along with a new
generation of research sat-
ellites— are described.

The Titanic
Revisited

Vol. 29:3, Fall 1986— The
second visit to the site, and
the first visit by Alvin and the
remote vehicle, Jason jr., is
described, and new findings
are reported. Other articles
address the radioactivity of
the Irish Sea, the growth of
U.S. aquaria, Japanese
ocean architecture, and the
collaboration of John Stein-
beck and Ed Ricketts.

The Great Barrier
Reef: Science &
Management

Vol. 29: 2, Summer 1986—
The Great Barrier Reef off
Australia's Pacific coast is
the world's largest coral reef
system. This comprehensive
special issue describes the
structure, evolution, life, and
management of this colorful
and complex system. Widely
useful to all with interests in
special ecosystems.

o o o

• The Arctic Ocean,

Vol. 29:1, Spring 1986 — An important issue on an active frontier

• The Titanic: Lost and Found,

Vol. 28:4, Winter 1985/86— The Jitanic's 1912 loss, and 1985 discovery.

• The Oceans and National Security,

Vol. 28:2, Summer 1 985 — The oceans from the viewpoint of the modern navy,
strategy, technology, weapons systems, and science.

• Marine Archaeology,

Vol. 28:1, Spring 1985 — History and science beneath the waves.

• The Exclusive Economic Zone,

Vol. 27:4, Winter 1984/85— Options for the U.S. EEZ.

• Deep-Sea Hot Springs and Cold Seeps,

Vol 27:3, Fall 1984 — A full report on vent science.

• El Nino,

Vol 27:2, Summer 1984 — An atmospheric phenomenon analyzed.

• Industry and the Oceans,

Vol. 27:1, Spring 1984

• Oceanography in China,

Vol. 26:4, Winter 1983/84

• Offshore Oil and Gas,

Vol. 26:3, Fall 1983

• Summer Issue,

1982, Vol. 25:2 — Coastal resource management, acoustic tomography, aqua-
culture, radioactive waste.

• Summer Issue,

1981, Vol. 24:2 — Aquatic plants, seabirds, oil and gas.

• The Oceans as Waste Space,

Vol. 24:1, Spring 1981.

• Senses of the Sea,

Vol. 23:3, Fall 1980.

• Summer Issue,

1980, Vol 23:2 — Plankton, El Nino and African fisheries, hot springs, Georges
Bank, and more.

• A Decade of Big Ocean Science,

Vol. 23:1, Spring 1980.

• Ocean Energy,

Vol. 22:4, Winter 1979/80.

• Sound in the Sea,

Vol. 20:2, Spring 1977 — The use of acoustics in navigation and oceanography

Send orders to:

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P.O. Box 6419
Syracuse, NY 13217

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