A Biodiversity Vision for the Galapagos Islands

G A L A P A G O S I S L A N D SA BIODIVERSITY VIS ION FOR THE

Based on an International Workshop of Conservation Biologists in Galapagos in May 1999

© Photo Heidi Snell

A BIODIVERSITY VISION FOR THE GALAPAGOS ISLANDS

Based on an International Workshop of Conservation Biologists

in Galapagos in May 1999

THE CHARLES DARWIN FOUNDATION

AND

THE WORLD WILDLIFE FUND Edited by R. Bensted-Smith A Spanish edition, translated by Isolda Rojas Lizana, is also available. The workshop participants were: Adsersen, Henning Allnutt, Tom Anderson, Dave Bensted-Smith, Robert Bostford, Loo W. Branch, George Bustamante, Rodrigo Causton, Charlotte Cifuentes, Miguel Cowie, Rob Cruz, Eliecer Christensen, Villy DeVries, Tjitte Dexter, Nick Dinerstein, Eric Dowler, Robert Eldredge, Lucius G. Espinoza, Fernando Ford, Glenn Gardener, Mark

Gaibor, Nikita Geller, Jonathan B. Gibbs, James Godbey, Maria Hickman, Cleveland Kakabadse, Yolanda Kendrick, Gary Marquet, Pablo Martinez, Priscilla McCosker, John Merlen, Godfrey Mooney, Harold Muñoz, Edgar Nafus, D.M. Paulay, Gustav Peck, Stewart Pellerano, Miguel Porter, Sanford Powell, George Reck, Günter

Rejmánek, Marcel Richmond, Robert Rivera, Fernando Ruiz, Ruth Elena Seddon, Mary Silva, Paul Smith, Cliff Snell, Howard Spurrier, Lauren Suárez, Luis Tye, Alan Ulloa, Robert Valle, Carlos Vargas, Hernán Veitch, Dick Wellington, Jerry Wikelski, Martin Witman, Jon

12 June 2002

How to cite this publication: Charles Darwin Foundation and World Wildlife Fund. 2002. A biodiversity vision

for the Galapagos Islands. Ed. R. Bensted-Smith. CDF, Puerto Ayora, Galapagos.

How to cite a chapter within this publication:

Tye, A., H.L. Snell, S.B. Peck and H. Adsersen. 2002. Outstanding terrestrial features of the Galapagos archipelago. In A Biodiversity vision for the Galapagos Islands. By Charles Darwin Foundation and World Wildlife Fund, Puerto Ayora, Galapagos.

Acknowledgements: This Vision is a collective effort and would not have been possible without the contributions of many people. World Wildlife Fund (WWF) and the Charles Darwin Foundation (CDF) would like to express our gratitude to each of the workshop participants and their home institutions. We would also like to thank the organizations whose support made the production of this Biodiversity Vision possible. In particular, we would like to thank:

• World Wildlife Fund-United States for sponsoring the Vision workshop;

• TAME Airlines for discounting air fares for participants; • The Government of Ecuador – and in particular the Galapagos

National Park Service - for their participation and support, including provision of the auditorium for the workshop;

• The United Nations Foundation and United National Development Program/Global Environment Facility for financial contributions to the printing of this document.

Many staff members and associates of WWF-US and the Charles Darwin Foundation were involved in conceiving and planning the workshop, designing the methodology, implementing the workshop, defining the Vision and writing the document. We cannot name all the people who have been involved in this long project, but would like to recognize in particular the contributions of the following people (in alphabetical order): Henning Adsersen, Tom Allnutt, Robert Bensted-Smith, Paulina Bermeo, George Branch, Rodrigo Bustamante, Charlotte Causton, Gonzalo Cerón, Eric Dinerstein, Graham Edgar, Isadora Espinoza, Glenn Ford, Maria Elena Godbey, Marie Louise Johnson, Priscilla Martinez, Stewart Peck, George Powell, Fernando Rivera, Sol Rojas, Marta Romoleroux, Poly Robayo, Franz Smith, Howard Snell, Lauren Spurrier, Alan Tye, Carlos Valle, Jerry Wellington, Jon Witman and Edwin Yanez.

General conclusion of the biodiversity vision analysis Galapagos stands at a crossroads: unlike other oceanic archipelagoes it still retains most of its original species, but ecological degradation is proceeding rapidly. With concerted conservation efforts, decisive policies and actions to address the root causes of the threats to Galapagos, the great majority of the islands’ unique biodiversity can be saved and even restored. Without such actions the degradation will accelerate. Certain development trends are in conflict with the conservation of Galapagos, especially the growth of three sources of pressure: human population, transport to and within the archipelago, and fishing. These trends are driving the depletion of populations of native species and the transformation of natural ecosystems by a rapidly increasing array of invasive alien species, from diseases and insects through to mammalian predators. Without radical and innovative measures to halt these trends and mitigate their impacts on the native flora and fauna, the processes of ecological change already under way will lead inevitably to loss of populations, extinction of species and disruption of ecosystems and evolutionary processes. In recent years the Government of Ecuador has embarked on some important initiatives, notably controlling migration to the islands, initiating a quarantine inspection system, increasing Park funding, obtaining large grants and a loan for conservation projects, and creating the Galapagos Marine Reserve, which excludes industrial fishing and is managed by the Park through a participatory management regime involving local stakeholders and partner institutions. The Government is also preparing a strategy for the sustainability of human presence in Galapagos, as well as a binding regional plan built on that strategy. It is essential that the strategy and the plan address the threats to biodiversity and guide Galapagos towards a sustainable future, in which a small, well educated, healthy human population co-exists with nature, uses resources sparingly and works constantly to control alien species. The people would have their own, distinctive way of life, appropriate to oceanic islands that evolved in isolation from man and are consequently so vulnerable to human presence. They would accept restrictions and responsibilities and enjoy to the full the privilege of living in one of the most special natural environments on Earth. We hope that the scientific analysis and projections presented here will persuade the Government of Ecuador to take well reasoned, responsible decisions that steer Galapagos away from irreversible loss of biodiversity and ecological degradation and towards sustainability and restoration.

TABLE OF CONTENTS

INTRODUCTORY SECTION CHAPTER 1 - PLANNING FOR THE ECOREGION R. Bensted-Smith, G. Powell and E. Dinerstein CHAPTER 2 – APPROACH TO PROJECTING THE FUTURE OF GALAPAGOS BIODIVERSITY H.L. Snell, G. Powell, A. Tye, R. Bensted-Smith, R.H. Bustamante and G.M. Branch. TERRESTRIAL SECTION CHAPTER 3 - OUTSTANDING TERRESTRIAL FEATURES OF THE GALAPAGOS ARCHIPELAGO A. Tye, H.L. Snell, S.B. Peck and H. Adsersen CHAPTER 4 - CONSERVATION CRITERIA FOR THE TERRESTRIAL BIOME A. Tye and H.L. Snell CHAPTER 5 - THE STATUS OF AND THREATS TO TERRESTRIAL BIODIVERSITY H.L. Snell, A. Tye, C.E. Causton and R. Bensted-Smith CHAPTER 6 – PROJECTIONS FOR THE FUTURE: A TERRESTRIAL BIODIVERSITY VISION H.L. Snell, A. Tye, C.E. Causton, G. Powell, E. Dinerstein, T. Allnutt and R. Bensted-Smith. MARINE SECTION CHAPTER 7 - OUTSTANDING MARINE FEATURES OF THE GALAPAGOS ARCHIPELAGO R.H. Bustamante, G.M. Wellington, G.M. Branch, G.J. Edgar, P. Martinez, F. Rivera, F. Smith & J.D.Witman. CHAPTER 8 - CONSERVATION CRITERIA FOR THE MARINE BIOME G.M. Branch, J.D. Witman, R. Bensted-Smith, R.H. Bustamante, G.M. Wellington, F. Smith & G.J. Edgar. CHAPTER 9 - THE STATUS OF AND THREATS TO MARINE BIODIVERSITY R.H. Bustamante, G.M. Branch, R. Bensted-Smith & G.J. Edgar CHAPTER 10 - PROJECTIONS FOR THE FUTURE: A MARINE BIODIVERSITY VISION R. Bensted-Smith, G.M. Branch, R.H. Bustamante, and G.M. Wellington CONCLUDING SECTION REUNITING MARINE AND TERRESTRIAL THEMES Chapter 11 : SUMMARY OF THE VISION AND CENTRAL ISSUES R. Bensted-Smith, T. Allnutt, G.M. Branch, R.H. Bustamante, C.E. Causton, E. Dinerstein, G. Powell, H.L. Snell, A. Tye, G.M. Wellington and J. Witman ANNEXES 1.1 List of workshop participants 1.2 Laws and policies 3.1 Physical setting of the Archipelago 4.1 IUCN Categories of Threatened Species 8.1 Proposed group of indicators of marine ecosystem composition and function

CDF/WWF Biodiversity Vision Chapter 1 – Planning for the Ecoregion

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CHAPTER 1 – PLANNING FOR THE ECOREGION Principal Authors – R. Bensted-Smith, G. Powell and E. Dinerstein 1.1 Introduction – “Back to Eden – one last chance” Galapagos, a name that to many is synonymous with pristine nature and scientific enlightenment, is also one of society’s greatest conservation opportunities and challenges. The archipelago is virtually unique on earth as a largely self-contained ecological system, or “ecoregion”, of high biological endemism, that could still be conserved as an example of how the world’s oceanic islands existed before modern humans intervened. The 128 islands that comprise the archipelago still retain 95% of their original, pre-human diversity of species, and important areas of modified habitat could eventually be restored, if radical steps are taken to control alien species and human activities. The surrounding seas, though less studied, also retain their diverse and dynamic biological communities and there appears to have been little irreversible loss of marine biodiversity. The unique island flora and fauna, which gave Charles Darwin enlightenment that changed the course of modern science, could continue to be available to enlighten future generations and inspire them to protect their world. On the other hand, current trends point towards continuing ecological degradation, with reduction of species abundance, loss of populations and eventually species extinctions. For 300 years after the relatively late discovery of the Islands in 1535, the use of natural resources was primarily extractive with little or no concern for sustainability. Colonization did not begin until the early 1800’s and remained at a low level until the 1960’s, when the Islands began to acquire a reputation as a tourist paradise. Since then, Ecuadorians from the mainland, attracted by economic opportunities, have arrived in large numbers. A 1998 census put the resident population at about 15,000 people, excluding tourists and other visitors, and annual population growth in the past decade has been 6%. People have brought with them, deliberately or accidentally, a range of exotic fauna and flora, whose spread is threatening unique biodiversity throughout the islands. Many residents derive their living directly or indirectly from tourism based upon the native flora and fauna. Others join the rapidly expanding fisheries sector, which is impacting the marine ecosystem through increased exploitation of reef fish, lobster and in the 1990’s sea cucumber and shark fin. In offshore waters the pressure has come from industrial and semi-industrial boats coming from the mainland and abroad in search of tuna, sharks and other pelagic species. Thus, Galapagos stands at a crossroads, with restoration still a possibility but degradation looming. This prompted the biologists attending a workshop on Galapagos biodiversity in May 1999 to coin the phrase “Back to Eden - one last chance”. The Ecuadorian Government has taken a number of important steps to establish biodiversity conservation as a fundamental objective for Galapagos, notably the inclusion of 96% of the 7,900 km2 land area in a National Park (established 1959) and the international commitment to protect Galapagos as a UNESCO World Heritage site and Biosphere Reserve. In 1998, following a change of the national Constitution, the Special Law for Galapagos restricted migration to the Islands, created a 130,000 km2 multiple-use Marine Reserve, increased conservation funding, and required that Galapagos be managed according to a plan that covers the whole region, including National Park, Marine Reserve and inhabited areas. In 2001 the World Heritage site was extended to include the Galapagos Marine Reserve. The present publication aims to support this national conservation effort, by analysing the current status and future prospects of the archipelago’s biological diversity, with a view to providing scientific guidance for the formulation of policies and plans for the ecoregion. It is based on research by resident and visiting scientists of the Charles Darwin Foundation (CDF) and on the proceedings of a scientific workshop, co-hosted by the World Wildlife Fund (WWF) and CDF, held at Puerto Ayora, Galapagos, in May 1999. The participants in the workshop were a select group of biologists with specialist knowledge of Galapagos and/or of key ecological processes operating in Galapagos. They are listed in Annex 1.1. This is a strictly biological analysis, which will need to be complemented by a social and economic analysis, focusing on areas of probable contradiction between conservation requirements and social aspirations. Given the insidious nature of the invasive species problem


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and the pressure on marine resources worldwide, the conciliation of conservation and development in Galapagos is a daunting challenge. Some would argue that is unachievable, but Ecuador is committed to demonstrating that it can be done and that commitment is at the heart of its planning for the archipelago. 1.2 Ecoregion-based Conservation Planning To undertake the biological analysis, WWF and CDF used a protocol of conservation planning known as Ecoregion-Based Conservation or ERBC (Margules et al. 2000). The procedure applies the principles of conservation biology to developing long-term plans that, when implemented, will conserve the full array of biodiversity and ecological processes native to the targeted region. The targeted region is generally an "ecoregion" or large unit of land or water that contains a distinct assemblage of natural biological communities sharing a large majority of species, dynamics, and environmental conditions. More specifically, the primary goals of ERBC are to: • Represent, in a system of protected areas, all native ecosystem types and serial stages

across their natural range of variation; • Maintain viable populations of all native species in natural patterns of abundance and

distribution; • Maintain ecological and evolutionary processes, such as disturbance regimes, hydrological

processes, nutrient cycles, and biotic interactions; • Maintain blocks of natural habitat large enough to be resilient to large-scale periodic

disturbances and long-term change (Noss 1991, Noss and Cooperrider 1994). The ERBC procedure requires the completion of an assessment of the current status of biodiversity in the ecoregion compared with pristine conditions (defined as prior to intervention by western civilisation), and the creation of a "vision" or definition of the minimum that is required in terms of areas, species, habitats, and ecological processes, to achieve the four conservation objectives in perpetuity. A biodiversity vision is considered an essential part of ERBC because it helps to move proposed actions beyond a business-as-usual approach to conservation. The Vision serves as a touchstone to ensure that the biologically and ecologically important features identified in the biological assessment remain the core conservation targets throughout the process of implementing conservation actions. To apply ERBC to the Galapagos Archipelago, CDF and WWF assembled a group of world-class biologists, who could apply their expertise, derived from studies of the Galapagos and the other major island systems of the world, to developing a systematic assessment of the status of biodiversity in the archipelago, assessing which components of the Galapagos flora and fauna are most threatened by human impact, and projecting what will happen over the long term if steps are not taken to mitigate the threats. The scientists were informed of Ecuador’s conservation objectives and policies for Galapagos and were charged with formulating a vision, based on their extensive experience and knowledge of the ecological processes at work, of what the archipelago would be like, in biological terms, in the year 2050, if it were managed optimally for conservation. Lastly, the scientists identified key issues that Ecuador will need to address, in order for such a biological vision to become a reality. Charles Darwin Research Station (CDRS) scientists and their collaborators prepared a summary of the distribution and status of Galapagos biodiversity, for consideration at the workshop. They have subsequently extended that baseline data, as well as ordering the results of the workshop deliberations and providing additional information, where appropriate. 1.3 Galapagos as a Model for Ecoregion-Based Conservation To maximise the likelihood of long-term success, an ecoregion plan should ideally have the following five characteristics:

a) Include the whole geographical area that could exert ecological influence on the biological community;

b) Be covered by a single administrative unit or a cluster of closely integrated administrative units, in which biodiversity conservation is already recognized as a significant management objective;


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c) Address threats at all scales, from specific, localised ones to more general, large-scale factors;

d) Be guided by a long-term “vision” and goals for biodiversity conservation and prescribe methodologies for monitoring progress towards those goals;

e) Consider the needs and aspirations of all the human communities with a significant interest in or influence on the biodiversity.

For any major area of biological diversity, the first two of these characteristics are generally in conflict. On the one hand, the validity of a plan that covers only part of an ecosystem will always be conditional on assumptions about what may happen in the rest of the ecosystem. On the other hand, any area large enough to be ecologically self-contained is likely to span several administrative regions and sectors, even multiple countries, for some of which biodiversity conservation may be a low priority. It is, of course, impossible for an ecoregion to be wholly self-contained; there will always be global external factors, such as global warming or the actions of the World Trade Organisation, which must be identified and taken into consideration. As far as possible, these should be limited to truly global factors, not local or regional ones which are best addressed within the planning process. The development of a meaningful biodiversity vision requires the availability of scientific information, which for many important ecosystems is scarce. Lastly, most ecoregions include numerous and diverse human communities, which means that the analysis of social and economic factors, to complement the biological analysis, is complicated and the implementation of the plan even more so. Consequently, in many regions ERBC planning has to overcome several obstacles in order to be effective. Scanning the world for ecologically self-contained regions in which the problems of administrative and social scale may be manageable, conservation is an established objective throughout the region, and scientific information is available, Galapagos stands out as the prime example. The ecological limits are relatively well defined, although the outer limit for wide-ranging marine organisms, such as pelagic fish or animals with planktonic larvae, cannot be precisely defined. For the terrestrial ecosystem, the predominant ecological interaction with the mainland is via man-made transport i.e. boats and planes. One environmental factor affecting Galapagos that is global in nature is climate, in particular the frequency and intensity of the El Niño phenomenon, which may be increasing due to global climate change. In administrative terms Galapagos consists of a single province of Ecuador, plus two protected areas, the Galapagos National Park (GNP) and the Galapagos Marine Reserve (GMR). The GNP covers 96% of the land mass, whilst the GMR extends 40 nautical miles from the so-called “baseline”, i.e. a line connecting the outermost points of land of the archipelago (Figure 1), and probably includes the ranges of most Galapagos marine life. An excessive number of government institutions govern the 26,356 ha inhabited area of Galapagos; the principal ones are the three Municipal Councils, a Provincial Council, the Provincial Governor, the National Galapagos Institute, known as INGALA, and the Ecuadorian Air Force, which occupies one island (Baltra). In addition there are provincial departments of several national ministries. This plethora of institutions, which is a consequence of the decision in 1973 to make Galapagos a province rather than a special territory, is an impediment to planning and management, but is a fraction of the number involved in most other ecoregions of the world. Significantly, this impediment has been greatly reduced by a constitutional amendment, enacted in 1995 and retained in modified form in Ecuador’s new constitution, adopted in 1998, and by the Special Law for Galapagos (SLG), also enacted in 1998. These legal instruments create obligatory mechanisms for province-wide planning, coordinated by INGALA. The preparation of the so-called Galapagos Regional Plan is being pushed along by INGALA during 2001/2. The Special Law also created the GMR and brought it under the administration of the Galapagos National Park Service (GNPS), thereby simplifying further the administrative set-up, as well as enhancing tremendously governmental commitment to marine biodiversity conservation.


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The same legislation establishes biodiversity conservation as one of the principal objectives of the whole province, the other being sustainable development. The SLG also includes important conservation considerations in the list of principles for all planning and management in Galapagos. However, although the SLG commits the Government to harmonizing conservation and development for the Islands’ residents, it does not set out a strategy for overcoming the potential contradictions between these twin objectives. Rather, it is a collection of conservation measures and social and economic privileges (see Bensted-Smith, 1998, for detailed comments on the SLG). This deficiency increases greatly the importance of the regional planning process for Galapagos, which, according to the SLG’s General Regulation, must include the development of a strategy for achieving sustainability of the presence of people in Galapagos. Further details of Ecuador’s laws, policies and objectives for Galapagos are included in Annex 1.2. Figure 1.1 Map of the Galapagos Marine Reserve (GMR) showing its platform (200 and 300 meter contour depths) and boundary (40 nautical miles from the baseline around the islands). In conclusion, Galapagos is not without its institutional complications but is administratively far simpler than most other ecoregions, as well as being biogeographically and ecologically a relatively well defined and well studied entity. Furthermore, the Government of Ecuador is committed to conservation of Galapagos biodiversity and to preparing a regional plan. Thus, Galapagos is a potential model ecoregion for the ERBC approach to planning. 1.4 The Purpose of This Document This document aims to elaborate Ecuador’s stated long-term goals for the conservation of the biodiversity and evolutionary processes of Galapagos into an overall vision for the biological state of the archipelago 50 years from now. This overall vision will be compared with projections based on an assumption of “business as usual”, that is to say without radical changes in conservation and development policies. The vision provides a benchmark and indicators, with which to measure progress in conserving the ecoregion, as well as a target to guide and motivate conservationists, planners and political leaders over the coming decades. The second chapter of this document describes the approach and methodology used to describe the status of Galapagos and projections for the future. Thereafter the document contains a terrestrial analysis (chapters 3-6) followed by a marine analysis (chapters 7-10), reflecting the fact that the two halves of the ecosystem have different biological characteristics, conservation indicators, status, threats and opportunities. The final chapter (11) aims to integrate and summarise the terrestrial and marine visions and identify some of the critical issues to be addressed, in order to progress towards the biodiversity vision. The document does not attempt to address socio-economic considerations; it presents only the biological perspective. Of course, government policy in general, and the Galapagos Regional Plan in particular, are concerned with both biodiversity conservation and sustainable

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development. Furthermore social and economic factors will be crucial in determining the strategies and plans of action for achieving conservation objectives. This biological analysis cannot on its own prescribe conservation plans, but it can help to define a national vision for the archipelago’s biodiversity, identify the key issues that need to be addressed in order to realize that vision, and suggest lines of action. These key issues should be the focus of a subsequent social and economic analysis. The combined biological and socio-economic analysis can form the basis for the strategy whereby Ecuador can seek to conserve the biological diversity and evolutionary processes of Galapagos, whilst enhancing the quality of life of the archipelago’s people. Specifically, we hope that this document will help regional planners by: • Translating Ecuador’s conservation goals for Galapagos into a vision of the long-term future

of the Islands’ biodiversity; • Providing an authoritative biological analysis as core information for the regional planning

process; • Giving concrete meaning to the term “sustainable” in the Galapagos context; • Serving as a tool for gauging the biodiversity implications of ideas proposed for inclusion in

the Galapagos Regional Plan; • Providing a yardstick, against which the success, in terms of biodiversity conservation, of the

Galapagos Regional Plan and its implementation can be assessed; • Providing a set of indicators to be used to measure the conservation of biodiversity and

evolutionary processes in Galapagos. REFERENCES Bensted-Smith, R. 1998. Comments on the Special Law for Galapagos (1998). Available on

the Charles Darwin Research Station website: www.darwinfoundation.org Margules, C.R. and Pressey R.L. 2000. Systematic Conservation Planning. Nature 405 (6783)

pp 243-53. Noss, R.F. 1991. Protecting Habitats and Biodiversity, Part 1: Guidelines for Regional Reserve

Systems. National Audubon Society, New York. Noss, R.F. and A.Y. Cooperrider. 1994. Saving Nature’s Legacy: Protecting and Restoring

Biodiversity. 380 pages. Island Press.

CDF/WWF Biodiversity Vision Chapter 2 – Approach to Developing Projections

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CHAPTER 2 – APPROACH TO PROJECTING THE FUTURE OF GALAPAGOS BIODIVERSITY Principal Authors – H.L. Snell, G. Powell, A. Tye, R. Bensted-Smith, R.H. Bustamante and G.M. Branch. 2.1 Introduction The scope of the development of a Biodiversity Vision for Galapagos is scientific in describing the biological and physical settings of the archipelago and the biological consequences of factors that are bringing about change in their natural state. Such factors are identified as best as is possible. Biological goals that should serve as measures of management success, are also presented. . However, the details of the management efforts required to achieve those goals are not within the scope of this volume. These may subsequently be developed based upon analyses of the goals identified herein, in the light of social and political processes within the Galapagos, Ecuador, and the world. The goals identified in the present document are based upon scientific analysis and are realistic in biological terms; how to reach them is what remains to be developed by experts in management. Because the Galapagos Archipelago consists of two major, interrelated, but distinct, ecoregions: a set of islands and their marine surroundings, it was necessary to structure the analysis process so that terrestrial and marine scientists could develop their assessments separately, but still integrate them as appropriate. Towards this end, the description of biological and physical settings, the discussion of status and threats, the projections of future biodiversity trends, and the articulation of management issues to be addressed, were implemented with parallel terrestrial and marine sessions. Constant dialogue between the two groups ensured their products were compatible for final integration. The Galapagos islands have been a focal point of conservation biology in Ecuador for many years. In that time a tight integration between research and management has evolved into the quasi-formal combination of evaluation (research and monitoring) and response (management). The goals are to respond to past, present, and future anthropogenic change in the biological diversity via restoration, mitigation, and prevention (Figures 2.1 and 2.2, Gibbs et al. 1999). Chapter 1 of this document reviewed the substantial, historical and on-going efforts by Ecuador to conserve the region's biodiversity. Annex 1.2 highlights national legislation pertinent to the conservation of the Galapagos. The review clearly demonstrates Ecuador's outstanding commitment to protecting this global resource. The chapters that follow reflect the dichotomy of terrestrial and marine analyses and projections by presenting them in separate chapters (3-6 terrestrial, 7-10 marine) with a final chapter (11) for integration. Chapter 3 reviews the outstanding terrestrial biological features of the Galapagos, particularly the baseline situation when the islands were discovered by western civilisation in 1535. Chapter 4 sets out future criteria for measuring the status of terrestrial biodiversity and for assessing changes in it. Chapter 5 presents the status of terrestrial biological diversity in 1999, and points out gaps in our knowledge of this. Chapter 6 develops projections of what is likely to happen to the terrestrial biological diversity of the Galapagos in the future given different levels of management effort. These projections are inevitably speculative in nature but represent the best judgement of the authors and other workshop participants, on the basis of what is known about the flora and fauna, the environment, the ecological processes at work, current trends and experience in other island ecosystems. Chapter 7 introduces the marine biome of the Galapagos Archipelago. Chapters 8, 9 and 10 follow the same structures as Chapters 4, 5 and 6, for the marine biome. Finally, Chapter 11 summarises the findings for both biomes, synthesises them into a combined Biodiversity Vision, and provides an overview of the issues that should be addressed for the vision to become a reality. This document tries to standardize the use of some potentially confusing terminology. Use of the three terms; species, infraspecific taxa, and distinct populations is necessary to encompass the range of variability potentially managed in the conservation biology of the Galapagos. “Species” refers to formally named species. “Infraspecific taxa” refers to named subspecies, races, or varieties. “Distinct populations” refers to apparently evolutionarily distinct populations that are not taxonomically recognized. Within different groups of organisms the


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degree to which these terms are used varies. In plants distinct populations are rarely addressed whereas in vertebrates populations are commonly emphasized. Some of these measures are loosely adapted from IUCN (1994). Furthermore, the term "indigenous" or "native" is defined as any species whose first occurrence in Galapagos is completely independent of anthropogenic activity; in this sense, "indigenous" and "native" include endemic species. An "endemic" species in an indigenous (native) species that is naturally found only in the Galapagos. In contrast, an "alien" or "introduced" species is defined as any species whose first occurrence in the Galapagos is a direct or indirect consequence of anthropogenic activity. Workshop Design:

The Biodiversity Vision Workshop brought together scientists and diversity managers experienced with the Galapagos or similar areas worldwide. This group met in May 1999 in Puerto Ayora, Isla Santa Cruz, Galapagos, Ecuador. In view of the uniqueness of the Galapagos situation, our approach to completing an ecoregional-based biological analysis and developing a vision for the future was modified from the standard Eco-Region Based Conservation (ERBC) procedures. For example, a complimentarity analysis of all vertebrate species by island demonstrates that all of the larger islands contribute unique endemic species. Even with the inclusion of the principal 10 islands, only 96% of the biodiversity of vertebrate species is represented. Thus, a representation analysis would call for the inclusion of at least all of these islands and would not be particularly helpful in setting conservation priorities.

Primary Goal:Preserve Biological Diversity in Natural State

Goals of Ecological Monitoring in Galapagos

Detect ChangesPotentialCurrent

Past

Identify Causesof Changes

AnthropogenicChanges

NaturalChanges

Evaluation

Response

Interactions

Prevent

Mitigate

Restore

Observe

Figure 2.1. Relationships among goals for conservation activities for the Galapagos. Solid lines and arrows indicate primary interactions and dotted lines are secondary. The monitoring program is primarily an evaluation activity and the subsequent management activities are responsive.

Objectives of Ecological Monitoring in Galapagos

Response

Prevent Potential Changes

A. Prevent Dispersal & Arrivalof ExoticsB. Protect Stable NaturalPopulat ionsC. Anticipate New DetrimentalHuman Activity

Mitigate CurrentPertubations

A. Control Exotics or EffectsB. Control DiseaseC. Recuperate or AccelerateGrowth of Declining PopulationsD. Control/Modify HumanActivity

Restore to BaselineA. Recuperat ion of NaturalPopulat ionsB. Eradication of Exot icsC. Cease Detrimental HumanActivity

Observe Natural Changes

Establish Baseline(Situat ion in 1534)

1. Organisms & Communities2. Ecological Relationships3. Evolutionary Processes

Establish Current SituationA. Deviations from baseline1. Distributions of Exotics2. Declines in Abundance3. Extinct ions4. Altered Ecology & Evolution

Monitor ChangeA. Historical:1. Rates & Distribution of Deviat ionsB. Current:1. Distribution & Abundance2. Reproduct ion & Mortality3. Individual Condition

Evaluate CausesA. Empirical - Within GalapagosB. Deduct ive - From Other Systems

Evaluation

Predict Future ChangeA. AnthropogenicB. Natural

Figure 2.2. Relationships among objectives of the ecological monitoring program. Solid lines indicate the usual progression between objectives. The dotted line indicates a less anticipated situation where changes of unknown cause can be predicted, perhaps due a cyclical temporal pattern.


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Similarly, a cursory analysis of threats revealed that invasive alien species threaten all islands regardless of their intact nature. Thus, for terrestrial components of the analysis we de-emphasized the usual focus on representation and intactness analyses and concentrated instead on developing a better understanding of overriding threats to biodiversity on the islands. For the marine components, ensuring representation of biogeographic regions and all habitats remained an important goal, but intactness was again played down, as all marine areas are under threat of overfishing. The workshop alternated between plenary sessions, for the presentation of general information, and small working groups, during which the information was discussed and specific recommendations were drafted. The proposed recommendations were then presented in plenary session by a representative from each working group and the floor was opened for discussion. Recommendations for conservation action were developed from the plenary sessions. In preparation for the workshop the CDRS prepared summaries of species distributions for 77 islands. The data were used to prepare basic support maps of terrestrial endemism (see Chapter 3). As another preparation for the workshop, WWF developed a general simulation model to predict the trajectory and dispersal of introduced species as a function of management decisions in the Galapagos archipelago. The model was designed to predict the relative probability of invasive species reaching specific islands under different human use scenarios. Data to drive the model were extracted from existing expertise on Galapagos invasives, research in other real and virtual island systems, and general findings from the field of invasion ecology. The plan was to present simulations of alien species invasions under different human use scenarios to assist participants make long-term predictions and recommendations for conservation actions. 2.2 Approach To Projecting The Future Of Terrestrial Biodiversity The projections in Chapter 6 are drawn from:

1. The baseline (what was Galapagos like prior to 1535), which provides both a benchmark and the basis for the ultimate long-term aspiration for biodiversity conservation;

2. The current status (what was Galapagos like in 1999); 3. What factors, reflected in patterns of the changes between 1535 and 1999, are

responsible for the difference? The projections are based upon:

1. A scenario that reflects “business as usual” – little change in the factors or in the management policies and efforts to mitigate the factors; known improvements in policies or efforts are taken into consideration.

2. An optimistic set of goals reflecting what is possible with large-scale changes in the

factors caused by great success in management efforts to mitigate the factors. This projection based on the assumption of optimal management is the basis for the biodiversity vision (see Chapter 6).

As already stated, the projections represent the best judgment of the authors and other workshop participants. Chapter 11 summarizes the vision and suggests what management issues need to be addressed to achieve it. Thus, Chapters 6 and 11 comprise a terrestrial “Biological Vision” of what the Galapagos could be in 50 years. Chapter 6 also highlights how the criteria of Chapter 4 can be used to monitor future change and assess our progress towards the vision. Establishing the baseline - what Galapagos was like in 1535 It can be difficult to establish a baseline from which changes in biological diversity can be measured. We are fortunate in the Galapagos because the ecosystem is relatively simple and the period of anthropogenic change is short. Our baseline for Galapagos is the status of


9

biological diversity prior to the first recorded visit by humans in 1535. It is possible that there were earlier visits, but there are no indications that earlier visits resulted in changes to the biological diversity. Observations made by the first visitors to Galapagos in 1535, of the presence or absence of all species on all islands, would comprise a hypothetical ideal baseline. Unfortunately, no visitors visited all islands, and none recorded all of the species observed. Historical information is piecemeal and often difficult to interpret. There are many shortcomings even with the limited observations that were made; for example, many observations lack voucher specimens and thus the accuracy of identifications can only be inferred. Therefore, the primary sources of information are direct data and inference. Direct data are drawn from the recorded observations mentioned above and physical records such as bones deposited in lava tubes by Barn Owls and pollen profiles in sedimentary deposits on the bottoms of ponds and lagoons. Unfortunately, there is no mineralized terrestrial fossil material known from Galapagos, due to the recent volcanic nature of the islands. Most of the direct data used in our analyses were drawn from the published literature. Inference about prehistoric biological diversity across the Galapagos includes extrapolation from observations of the present status and distribution. In the vast majority of cases, current or historically recorded presence, combined with knowledge of a species’ dispersal mechanisms, indicates presence for the baseline. Use of these data permits identification of presumed native and presumed introduced species, although there remains a group of species for which their origin (native or anthropogenic) is, and probably will remain, unknown. Establishing current status — what is Galapagos like today? The data used to establish the current status of the biological diversity of Galapagos come from published accounts, museum collections and anecdotal observations, augmented by recent fieldwork. Knowledge about the actual distribution of biological diversity is constantly growing, with frequent newly-documented occurrences. Up to about 1980, significant numbers of species were being added to the floral and faunal lists of many islands and one of the best statistical predictors of the number of native (or presumed native) species known to occur on an island was the number of scientific visits made to the island. However, since then, additions of native species have been few, and the number of scientific visits to an island no longer predicts the number of native species known to occur there (H. Adsersen, personal communication). As of 1994, reasonably complete lists of the vascular flora and vertebrate fauna had been compiled for 77 of the Galapagos islands. These data form the bulk of the material analyzed in chapters 3 and 5. The islands missing from this data set represent less than 0.01% of the total land area of the archipelago. A more significant gap in the data is represented by the “lower” plants and terrestrial invertebrates, including especially the soil fauna and flora, whose distributions have been comparatively understudied to date. Probably at least 10% of these groups remains to be discovered, and even more in the case of the soil organisms. Nature of changes, and identification of factors causing change The nature of changes is measured in several ways, including abundance, extinction or disappearance (for populations and species), extent (for habitats and communities), evolutionary shift (within a population or species), change in species composition (for communities), and alteration of habitat (for species and communities). We recognize that there are many other means by which change could be described. Our choice is based upon the available data, the feasibility of using the various criteria available, and the goals of this project. Changes in biological diversity and its patterns of distribution across the Galapagos are identified as differences between the current and baseline situations. Changes could be natural or anthropogenic, and in some cases it can be difficult to determine whether an observed change in biodiversity has a natural or an anthropogenic cause; in such cases we present justifications for the assignments. To determine the factors responsible for change, we examine the observed patterns of change in the light of factors known to be acting in Galapagos, or known to be responsible for such change in other areas over a comparable period of time (465 years). The potential causes of natural change include climatic fluctuations and long-term trends, volcanism, ecological succession, competition, predation and dispersal. The potential anthropogenic causes of


10

change include the effects of alien species, introduction of diseases, extractive use of resources and habitat alteration. 2.3 Approach to Projecting the Future of Marine Biodiversity The Galapagos marine and terrestrial biomes interact in many ways. Indeed, most Galapagos tourism depends on wildlife that lives part of the time on land but feeds in the sea, for example sea lions, fur seals, marine iguanas (the world’s only sea-going lizard), sea turtles and the abundance of seabirds and shorebirds. However, the threats in the marine environment and in the terrestrial environment are quite different. Marine biodiversity and ecosystem functioning are threatened principally by increasing exploitation of key species, This pressure could interact with the effects of climate change and introduced species. In addition, the marine environment has until very recently had neither effective protection nor effective management. By contrast, 96% of the terrestrial environment has been protected for 40 years, and in most of that area there is no extractive use whatsoever, but the flora and fauna face the overwhelming threat of invasive species, which may eat them, compete with them, carry diseases or transform their habitat. Furthermore, whereas the marine biome has quite a high diversity of species, the terrestrial biome has the depauperate flora and fauna typical of remote islands. Consequently, the approach to projecting the possible future of the marine biodiversity of Galapagos and of developing a marine biodiversity vision for the archipelago, differs slightly between the marine and terrestrial environments. Consideration of the interactions that affect coastal species that depend on both environments can be added, once the results for marine and terrestrial biodiversity have been derived. The marine analysis adopts the same basic approach of assessing the current status, threats and opportunities, then making projections based on various assumptions about the underlying factors. However, unlike the terrestrial environment, there is little evidence of irreversible ecological change in the marine environment of Galapagos, so there is less of a contrast between an idealistic vision and a realistic but imperfect one. There is abundant scientific evidence that over-exploited and heavily impacted marine ecosystems do recover, given sufficient protection and time. Most exploited marine species quickly recover when fishing ceases. In the case of Galapagos marine ecosystems, where negative impacts have not yet reached critical levels, there is great opportunity for recovery and restoration of affected species and habitats. Consequently, the description of a biodiversity vision is relatively straightforward. When developing the marine vision, the assumptions to be made about management of the marine environment mainly concern the alleviation of direct human impacts. Assumptions for “business as usual” projections are harder to define, because the management of the Galapagos marine environment is in a period of rapid change. After centuries of unrestrained exploitation, starting with the 18th century whalers, the marine environment of Galapagos at last obtained meaningful legal protection under the Special Law for Galapagos (SLG, 1998). The law created a protected area of 130,000 km2, banned industrial fishing within it, instituted a participatory process for planning and management, and required the development of a Management Plan with a zoning scheme. On the other hand, the past few years have also seen a rapid increase in fishing effort and persistent demands to exploit a wider range of species with high economic value. Furthermore, the industrial fishing sector is seeking the repeal of the marine conservation provisions of the SLG. In the light of this fast-changing situation, three marine projections were generated: the ideal vision, business-as-usual based on an assumption of only partial implementation of laws and plans and continuing pressures to expand fisheries, and an intermediate projection based on an assumption of markedly improved but not ideal management and control. Lastly, a distinctive feature of the approach to projecting marine biodiversity conservation is the recognition that the Galapagos marine ecosystem is highly dynamic and variable (Chapter 7). Thus, conservation goals are oriented towards ensuring the resilience of the ecosystem in the face of anthropogenic pressures and environmental variability, either local or global, as well as preserving species and populations.


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REFERENCES Gibbs, J. P., H. L. Snell, and C.E. Causton. 1999. Effective monitoring for adaptive wildlife

management: lessons from the Galápagos Islands. Journal of Wildlife Management 63(4):1055-1065.

IUCN. 1994. IUCN red list categories. IUCN, Gland.

CDF/WWF Biodiversity Vision Chapter 3 – Outstanding Terrestrial Features of the Galapagos

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CHAPTER 3 – OUTSTANDING TERRESTRIAL FEATURES OF THE GALAPAGOS ARCHIPELAGO Principal authors: A. Tye, H.L. Snell, S.B. Peck and H. Adsersen

The Galapagos Islands still represent a magnificent showcase of biodiversity due to their late human colonization. In no other large oceanic archipelago has human influence been so limited in space and time. The islands thus constitute a nearly unique example of how ecological, evolutionary and biogeographic processes shape the flora and fauna of single islands and an entire archipelago. The archipelago form itself promotes evolutionary change, with many different degrees of geographical and consequent genetic isolation. Isolation allows closely related populations to follow separate evolutionary tracks and leads to speciation and diversification. The universally known Darwin’s finches and giant tortoises are but two examples of the clarity with which Galapagos organisms demonstrate evolutionary processes. Many other components of the biota have evolved in isolation into organisms that are present nowhere else on earth, including, for example, endemic snails, insects, cacti, trees, rodents and iguanas. The Galapagos Islands are also important as the site where Darwin collected specimens reportedly crucial in developing his theory of evolution by natural selection. Evolutionary concepts have transformed human understanding of the natural world, and the implications of the evolutionary origins of humans have profoundly changed our ideas about our place in the natural world and our attitude towards its other inhabitants.

PHYSICAL SETTING

There are approximately 128 named islands recorded within the Galapagos (Annex 3.1). The total number of islands varies depending on how an island is defined and as more exploration within the archipelago is completed. This analysis deals with the 124 islands known or thought to be vegetated by at least one species of terrestrial plant other than mangroves and permanently isolated (Annex 3.1). These islands range in altitude from 2 to nearly 1700 m, in area from <0.001 to 4600 km2, and in isolation from 0.003 to 175 km (Figure 3.2 and Annex 3.1). Island size and isolation increase with distance to the nearest larger neighbor, but are not correlated with distance from the center of the archipelago. However, the most distant islands are intermediate in size and no large islands are more than 150 km from the center (Figure 3.2). The greatest extent of the archipelago is 431 km NNW from the southeastern corner of Española Island to the northwestern corner of Darwin Island.

Figure 3.2. Relationship of island size to isolation. Larger islands are significantly more isolated from larger neighbors than smaller islands (p < 0.01). Some, but not all, of this pattern could result from how proximate isolation is measured (see Annex 3.1, note 9). There is no consistent relationship of island size and distance from the center of the archipelago.

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AGE AND GEOLOGICAL HISTORY

As are all other oceanic archipelagos of the world, the Galapagos are of volcanic origin. Their source is a magma plume resulting in a “hotspot” under the crust of the earth. As the plates move over this hotspot, eruptions produce lava that can build up to form islands that drift eastwards on the Nazca Plate, away from the plume. The patterns of plate-movement that have produced the Galapagos are complex but, in general, the younger large islands form the western parts of the archipelago and the eastern islands are older. Some of the smaller central islets, probably formed by “parasitic” eruptive events, may be very young, possibly less than 10,000 years (D. Geist, personal communication), and even the oldest islands carry bare lava flows, evidence of fairly recent eruptions. Further to the east, towards the South American continent, a series of submerged seamounts may represent even older islands originating from the Galapagos Plume, which may have been above sea level within the last 10 million years, thereby extending the age of the archipelago considerably. The oldest of the current islands appear to have originated within the last 3–6 million years (Geist 1996). The youngest of the large islands, Fernandina and Isabela, are less than 1 million years old and Fernandina could even be as young as 60,000 years (Geist 1996).

Compared with most other oceanic archipelagos, the Galapagos is very young. The oceanic origin and youth of Galapagos have important consequences for its biological diversity. First, compared with a continent, or an older archipelago, relatively few species are present. Every terrestrial species has to arrive across an ocean barrier, survive once arrived, and establish a viable population, or has to evolve from a species that has crossed that barrier, survived and become established. There has also been little time for species to arrive and accumulate. Second, there has been comparatively little time for evolution of species, and development of species-rich communities. Plant communities are very simple, both structurally and in terms of species diversity. There has also been little time for the development of soils and species-rich communities.

CLIMATIC SETTING AND HISTORY

The climate of the Galapagos is atypical for a tropical oceanic archipelago. The varying presence of cold water brought north from the southern ocean by the Peru (Humbolt) Current cools and drys the Galapagos during much of the year. When those currents weaken and warmer more typically tropical waters from the north surround the islands the warmer rainy season occurs. The general pattern of a warmer, rainy season from January through April or May and a cooler, drier, garua season from June or July through October or November (occasionally into December) can be abruptly altered during El Niño events (Snell and Rea 1999). El Niño events result from a complex interaction of variation in the trade winds and the distribution of masses of warm water in the Pacific. The result is that warm waters remain around the Galapagos for long periods of time and intense and prolonged rains occur. These patterns make annual and seasonal variation in rainfall and temperatures within the Galapagos extreme (Figure 3.3). El Niño events have become more intense and frequent during the last 100 years with a peak frequency and intensity during the last 20 years (Snell and Rea 1999).

Raised bogs show that the climate of Galapagos has been humid in the highlands for the past 5000 years, but no older bogs are known (Colinvaux 1984). The only old lake basin known in Galapagos, at El Junco on San Cristóbal, contains sediment accumulated over at least 50,000 years, and its lake has not dried during the past 10,000 years, since the end of the last ice age (Colinvaux 1984). Before that, it was dry, with air-weathered sediment, but there is evidence of an even earlier wet period, dating from at least 48,000 years BP (Colinvaux 1984). These data indicate that the lake, and the high precipitation in the highlands which is necessary for its maintenance, are interglacial phenomena, with drier periods during the glaciations. This suggests that Galapagos would have passed through a dry period from 25,000 to 15,000 years ago, during which time humid habitats would have been at least much more restricted than at present, and perhaps completely absent from some islands that currently have them. The effect on evolution in the Galapagos would be that species adapted to dry climates would have had a longer period for speciation than humid-adapted species.


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BIOLOGICAL DIVERSITY

In this section we review the original status of the biodiversity of Galapagos: its key features and status at the time of discovery in 1535.

Habitat diversity and plant communities

Galapagos terrestrial habitats are defined to a great extent by their plant communities and rainfall patterns. Rainfall patterns are influenced by topography, aspect and position within the archipelago and the plant communities respond to all of these factors plus the geological age of the site. Plants determine the structure of the environment, with the vegetation structure superimposed upon the topography. There are four universally recognized vegetation zones, which occur throughout the archipelago: Littoral, Arid (in fact technically semi-arid), Transition and Humid. The Humid Zone is often sub-divided, with the divisions (including Scalesia, Miconia, Brown, Fern-sedge) varying from island to island. Although not normally sub-divided, the Arid Zone equally deserves to be, with a lower scrub zone and upper woodland zone, and the scrub zone being perhaps the most varied zone of Galapagos in terms of local community dominant species. The vegetation zones are a result of the Galapagos climate, where prevailing winds come from the southeast for most of the year. These produce higher

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precipitation on the southern slopes. Precipitation is also greater in the higher regions, due to orogenic cloud formation and condensation.

Within the major vegetation zones, Galapagos plant communities further define the environments available for its faunal inhabitants. Table 3.1 presents a list of well-defined plant communities. The diversity of communities represented is striking for such a small land area, in such a young geological setting.

Table 3.1. Galapagos plant communities, their dominant species, abundance and representation of (island) sub-communities in areas free of human impact.

Community Dominant species1 Abundance2 Subcommunities protected3

Littoral Zone Mangrove Avicennia nitida

Laguncularia racemosa Rhizophora mangle Conocarpus erecta

2 all

Sandy beach Ipomoea pes-caprae Scaevola plumierii

2 all

Dunes Mollugo spp., Amaranthus sclerantoides Polygala spp., Tiquilia spp.

3 some

Salt marsh Salicornia fruticosa Batis maritima

3 all

Brackish lagoons Ruppia maritima, Eleocharis spp. 3 some

Arid Zone Opuntia-Scalesia open scrub

Opuntia and Scalesia spp 1 some

Saltbush Cryptocarpus pyriformis 1 all Muyuyo scrub Cordia lutea 2 all Mesquite Prosopis juliflora 2 all Bursera dry woodland

Bursera graveolens B. malacophylla

1 all

Croton scrub Croton scouleri 1 some Cotton scrub Gossypium darwinii 2 all Dry highland scrub Macraea laricifolia 3 some Dry highland grassland

Pennisetum pauperum and other Poaceae

3 some

Transition Zone Pisonia woodland Pisonia floribunda 2 some Guayabillo woodland

Psidium galapageium 1 some

Scalesia-Guayabillo forest

Psidium galapageium with Scalesia tree spp

1 some

Humid Zone Scalesia Zone Scalesia tree spp, Psychotria spp,

Alternanthera halimifolia 2 none

Brown Zone Zanthoxylum fagara with Frullania 3 none Miconia Zone Miconia robinsoniana 3 none Acnistus scrub Iochroma elliptica 3 some Broad-leaved Croton scrub

Croton scouleri 2 some

Fern brake Pteridium aquilinum and other Pteridophyta

2 some

Pampa Cyperaceae and herbs 3 some Tree-fern groves Cyathea weatherbyana 3 none Fens 3 Vertical bogs Sphagnum spp. 3 some


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Pioneer series Ipomoea on lava Ipomoea habeliana 2 some Early lava pioneers Brachycereus nesioticus

Mollugo spp. 2 some

Late lava pioneers Jasminocereus thouarsii Darwiniothamnus spp Scalesia spp

2 some

Fumaroles Pteridophyta 3 some Ash plains Tiquilia spp. 2 some Tufa and scoria scrub

Macraea laricifolia Lecocarpus spp

2 some

Aquatic habitats Streams no widespread dominants 3 some Springs Potamogeton spp. 3 some Temporary ponds Algae 3 some Lakes Eleocharis spp 3 none 1 Under “Dominant species”, co-dominants normally occurring together are given on the same line, while

communities often dominated by one or other are given separate lines.

2 Relative representation is indicated under “Abundance” on a scale: 1 dominant over large areas of many islands; 2 dominant over substantial areas on few islands or smaller areas on many; 3 restricted to small areas. Note that these numbers refer to the community type, not to the abundance of dominant species (e.g. although Opuntia-Scalesia open scrub receives “1”, individual Scalesia and Opuntia species may be very rare).

3 Most of these plant communities have distinct sub-communities in different islands, each with its own species composition, in many cases including species endemic to that particular island. For example, the lava pioneer series have among their major components genera such as Mollugo and Scalesia, which are among the most striking examples of radiative evolution in Galapagos, such that the community on each island is structurally similar but made up of different representatives of the same genera. To complement the abundance indicator, the last column in the table indicates our provisional assessment of which of the communities currently have all, some or none of their sub-communities represented in areas isolated from anthropogenic impacts.

Species diversity and endemism

A separate paper (Snell et al. in prep) demonstrates that, on a global scale, the flora and fauna of Galapagos have relatively few species (low diversity). This is partly a result of their being an oceanic archipelago, partly a result of their youth relative to other such archipelagos, and partially due to habitat characteristics. However, Galapagos is famous for its “endemic” species.

Do the Galapagos warrant special consideration in efforts to conserve global patterns of biological diversity? The analyses mentioned above suggest that the Galapagos are not a striking “hotspot” of biological diversity, except perhaps for endemic vertebrates and possibly plants – given the young age of the archipelago. However, in later sections of this volume we compare the Galapagos with global patterns of extinction. The combination of those comparisons with these results highlight the significance of Galapagos to global biological diversity in today’s world. The Galapagos are a rare remnant of a prehistoric pattern of global biological diversity where great proportions of the worlds distinctive and often bizarre species occurred on islands. Man has destroyed much of that pattern. The biological diversity of the Galapagos is one of the best examples of that pattern because it remains. Few, if any, other options exist for preserving intact biotas of archipelagos. This is why the Galapagos warrants special attention – it is one of our last chances.

Evolution of one endemic species from a colonizing ancestor is one type of evolution (phyletic or linear) that is commonly found on oceanic islands. However, evolution often involves divergence, with one original colonizing species giving rise to several endemic species. Darwin’s finches, the giant tortoises and many other groups are excellent examples of such “evolutionary radiation”, where one original species has evolved into a whole range of different forms in their island isolation. The most spectacular example of such radiation has been in the snails of the family Bulimulidae, which has (or had before current extirpations began) more endemic species per island than even the famous finches.


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The native flora of Galapagos includes 560 species of vascular plants, and more than 600 taxa, including subspecies etc. This total includes a group of some 60 questionable species, principally pan-tropical weeds, for which there is uncertainty as to whether they arrived naturally or were brought (most likely inadvertently) by early human visitors, in the early years before much botanical work had been carried out. The totals include 180 endemic species and over 200 endemic taxa. Plant endemicity is therefore 32% at the species level, and is higher at the level of infraspecific taxa.Endemism and radiation have taken place in the plants of Galapagos to at least the same degree as in the animals. The endemic genus Scalesia of the family Asteraceae (Compositae) contains 15 species and a total of 19 generally accepted taxa, all of which are thought to have evolved from a single ancestral colonizing species. Plant endemism and radiation are not limited to the Scalesias. Some 53 families of vascular plants have evolved one or more endemic species in Galapagos. Of the 560 or so native Galapagos vascular plant species, about 180 (32%) are endemic (Lawesson et al. 1987). If one excludes the 60 doubtfully native species, the percentage of endemics is even higher (43%; see Porter 1983). Among the endemic plants are seven endemic genera: Darwiniothamnus, Lecocarpus, Macraea and Scalesia (all in the Asteraceae), Brachycereus and Jasminocereus (Cactaceae) and Sicyocaulis (Cucurbitaceae). Endemism is highest in the flowering plants, with the ferns and allies (Pteridophyta), mosses and liverworts (Bryophyta) and Fungi having much lower levels of endemism, probably due to their small spores and efficient dispersal, which reduce the isolation necessary for the evolution of new species. Endemism in the lichens of the Littoral Zone is quite high (Weber & Gradstein 1984), while the levels of endemism for lichens of other zones are low.

Examples of such radiation, which have produced at least three endemic taxa, are seen in 11 families and 19 genera of Galapagos vascular plants: Amaranthaceae (Alternanthera 8 taxa; Froelichia 5), Asteraceae (Darwiniothamnus 4; Lecocarpus 3; Scalesia 19), Boraginaceae (Cordia 3; Tiquilia 3), Cactaceae (Jasminocereus 3; Opuntia two independent radiations from separate colonizing events, resulting in 5 and 9 taxa each), Euphorbiaceae (Acalypha 6; Chamaesyce 9; Croton 4), Lamiaceae (Salvia 3), Molluginaceae (Mollugo 9), Piperaceae (Peperomia 4), Poaceae (Aristida 4; Paspalum 3), Polygalaceae (Polygala 5), Rubiaceae (Borreria 6). A lesser level of radiation has produced two species in each of 10 other genera, while 53 additional genera have a single endemic species. In some cases, evolution of endemic species has occurred from more than one colonization in a genus; for example, in four genera, two separate colonizations have each produced endemic species, while in Cyperus and Ipomoea three separate colonizations by species in each genus have each evolved into endemic species. In four genera (Cordia, Opuntia, Verbena and Alternanthera), two or more colonizations (five in the case of Alternanthera) have given rise to at least one endemic taxon each, and in the case of Opuntia the two introductions have each produced a large radiative group.

Plant endemism is higher in the lowland, more arid habitats of the islands, where 67% of the endemic vascular species are found, than in the more humid, highland vegetation types (29%), reflecting the longer period of time during which evolution can have proceeded in such habitats (see climatic history, above). Further, almost all examples of radiation are found in or at least commenced in the arid zone, with the only exceptions being Darwiniothamnus and Peperomia. The remaining 4% of endemics are littoral zone plants; very few littoral-zone plants are endemic, reflecting their ease of dispersal by ocean currents.

Galapagos is a typical oceanic archipelago in another respect: taxonomic disharmony. The filtering of species by the barriers to arrival and establishment results in an unusual selection of species reaching the islands, compared with the range of species available in the continental source areas. For plants, species that have wind- or bird-dispersed seeds are more likely to reach oceanic islands, so plant families with these characteristics tend to be over-represented compared with their nearest mainland areas. Plant families that are common on islands, including Galapagos include the Asteraceae, with their light, wind-borne seeds. Chance also plays a role: there may be many species representing a family on an archipelago, but all endemic and derived by radiative evolution from a single colonizing event. In Galapagos, this could be the case for the Polygalaceae (5 species) and Piperaceae (4 species), among others. Selection at the establishment phase is also important, with families with pioneer characteristics and that can survive on relatively new, soil-less lava (such as Cactaceae, Molluginaceae), being over-represented in Galapagos. These groups contain many of the Galapagos endemics. Taxonomic disharmony contributes to the vulnerability of the Galapagos biota, in that introduced


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species are often from families which would not naturally have been able to arrive and establish in the islands, which can thus produce profound changes in the character of the flora and fauna.

In terms of numbers of species (c. 1900 native species), invertebrates represent most of the natural terrestrial biodiversity of Galapagos (Peck 1997), although, as with other groups of animals and plants, Galapagos is depauperate relative to continental source areas. As in plants and vertebrates, some insect Orders are completely absent from Galapagos, such as the mayflies (Ephemeroptera). This “disharmony” is also evident within orders that are present, such as beetles (Coleoptera) and moths (Lepidoptera), where half or more of the Neotropical Families are not represented in Galapagos (Peck 1996). Invertebrates are involved in important ecological processes such as decomposition, pollination and spread of pathogens, but little is known in detail about these processes in Galapagos, with most conclusions being based on inference from morphology, and studies elsewhere.

The non-insect terrestrial invertebrates represent 71 families, 117 genera and 386 species, of which 363 are presumed native, including 193 endemic, and 23 introduced species (Baert 2000). The largest group is the Acarina, and the number of these may double with further taxonomic study (Schatz 1991). Levels of endemicity are unknown but probably around 50%. The land snail fauna of Galapagos is very diverse, with 83 indigenous species, 80 of which are endemic. The Bulimulidae is the most species-rich land snail family in Galapagos, represented by 65 species, all endemic (Chambers 1991).

An initial taxonomic inventory of insects is nearing completion, with 1822 species now known, of which 1530 are native, including 712 (47%) endemic, and the remaining 292 introduced (Peck 1996, Peck et al. 1998). Galapagos is not very diverse by comparison with sites in the lowland wet tropics, but is comparatively rich considering its environmental youth, isolation, and seasonally harsh climates. Galapagos is the world’s last little-altered insect ecosystem where we can identify patterns that existed before homogenization by introduced species of “weedy” insects.

There are 23 endemic insect genera, suggesting early arrival and long separation from mainland ancestors. The older islands such as Española, San Cristóbal, and Santa Fé appear to have more endemics. At least 50 genera of insects have undergone speciation, especially flightless ones such as carabid and darkling beetles (Stomion, Ammophorus and Blapstinus), and issid bugs (Peck 1996). These species have evolved from the first colonizing species of the lowlands, expanding their range and habitats to occupy the higher, humid zones. Examples of parapatric speciation include the litter dwelling Pterostichus beetles (Desender et al. 1992), lycosid spiders on Santa Cruz (Baert personal comm.) and cave dwelling insects (Peck 1996), as well as the Bulimulidae (Coppois 1984).

The highest diversity of terrestrial arthropods is found in the arid lowlands (Peck, Baert personal comm.), the largest zone in the archipelago. Zonation is less marked than with plants, perhaps because of better dispersal capacities, and distribution ultimately depends on feeding and habitat requirements, e.g. monophagous herbivores are restricted to areas where their host plant is found.

Patterns of diversity and endemism are simpler in vertebrates, primarily due to the reduced number of taxa (approximately 117 taxa, overall endemism 59%). As was demonstrated in plants and invertebrates, the less mobile groups have higher percentages of endemism (reptiles & terrestrial mammals > terrestrial birds > marine birds & marine mammals; Table 3.2). In many cases the taxonomy of Galápagos vertebrates is in dispute. When referring to vertebrates thoughout this volume we use an “evolutionary species” concept and treat distinct, isolated populations with independent evolutionary trajectories as components of biodiversity.


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Table 3.2. Species Richness and Endemism of Vertebrates.

Group of Organisms Total Taxa % Endemism

Reptiles 40 100

Birds 58 52 Marine 19 26

Aquatic/shore 13 23

Terrestrial 26 84 Mammals 16 88

Terrestrial 12 100

Marine (not cetaceans) 2 50 Bats 2 50

Patterns of distribution

Larger islands harbor more species than smaller islands, but the exponent of the relationship is less than one, meaning that an island twice as large as another is not likely to have twice as many species. The number of non-endemic native species increases more rapidly with island size than the number of endemics, resulting in a pattern where larger islands have a lower percentage of endemic species in their native diversity than smaller islands (Figure 3.4).

Larger islands also contain a greater percentage of the total diversity of Galapagos, both in native species and endemic species. However, there is an interesting interaction with the patterns of variation of native species of plants and vertebrates, endemic species of plants and vertebrates, and island size. The percentage of Galápagos diversity in endemic species of plants changes with the size of islands in a manner statistically indistinguishable from that of vertebrates. However the percentage of Galápagos diversity in native species of vertebrates is much greater than that of plants on smaller islands but nearly the same on larger islands (Figure 3.5).

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Figure 3.4. Galápagos Patterns of Species Richness. Data of island size from Table 2A.1. Regressions of Native and endemic species richness and island size significant (p<0.01). The regression of endemic species as a percentage of native species richness is not significant, although the “step” at 65% and 0.01 km2 is.


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Genetic Diversity

In the few groups that have been examined, Galapagos endemic organisms show very little genetic variability, whether within or between populations, or even between related species (Rick & Fobes 1975, Elisens 1989, D. Anderson pers. comm.). This is a trait common in radiative groups on oceanic islands (e.g. see Lowrey & Crawford 1985, Crawford et al. 1987), and reflects their recent origin from the arrival of a few individuals on the islands, which carried a very small proportion of the genetic variation present in their parent mainland populations.

Although other studies are now in progress, only one insect group has so far been examined for genetic diversity by allozyme techniques: the Stomion group of flightless darkling beetles (Finston & Peck 1995). Intriguingly, in this group, genetic variability patterns of genetic species do not match those of morphological characters for morphologically defined species.

Figure 3.5. Relative Distribution of Galápagos Diversity and Island Size. See text for interpreation of patterns.

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Evolutionary processes

The evolutionary processes that have led to the diversity of endemic species in Galapagos are well understood in very general terms, but have not been investigated in detail for most of the Galapagos flora and fauna. The present islands are between 3 and 6 million years old, and evolution must have been rapid to produce such diversity in such a short time. Radiation and extensive speciation is prevalent on very isolated and old archipelagos such as the Hawaiian Islands. However, radiation is an exceptional event, rather than the rule, in Galapagos organisms. Most plant, invertebrate and vertebrate colonists that have been present for long enough have speciated, but comparatively few have radiated.

The presumed closest relatives of the endemic plants of Galapagos are found in the following regions (Porter 1983): Andes and tropical Americas (79%), pantropical (8%), north and central America (7%), Caribbean (5%), rest of south America (1%). Most of the ancestors of the endemics were probably transported to the islands as seeds by birds (60%), wind (32%) and ocean currents (8%).

The presumed ancestral species of only about 15% of the Galapagos endemic vascular plants occur at present in the islands, indicating that for the other 85%, the entire colonizing population has evolved into taxa different from the colonizing species, or that the colonizer differentiated and then the original form became extinct. Most of the groups that have shown a lot of radiation do not have their presumed ancestor still present in the islands. The taxa that make up the groups that have radiated most, often have very restricted distributions, such as only one island, or part of an island. In contrast, endemic species that are not members of a radiative group, but which have been produced by linear rather than radiative evolution, tend to be more widely spread in the islands. Interestingly, the endemic plants of Galapagos tend to be found on more islands than the non-endemic native species.

The species per genus ratio in terrestrial invertebrates is less than 2 (a pattern that may be influenced largely by insects), confirming the comparatively low occurrence of radiation. A few groups of insects and other terrestrial invertebrates have undergone extensive radiative speciation. Examples are a few genera of ground dwelling beetles, crickets, plant-sucking bugs, and the endemic gastropod family Bulimulidae. Complexes of closely related species have resulted but, for insects, there is little evidence that they have developed different niches in different habitats or moved to different foods or feeding types. The most spectacular example of a habitat shift to a new environment is the part of the terrestrial invertebrate fauna that has become adapted to soil and cave environments. These have lost their eyes, and sometimes the thick cuticle, and gained longer legs or antennae. This process has not been accompanied by the production of many closely related species in these groups, so is adaptation without significant radiation.

The presumed closest relatives of the endemic insects are in the following regions: 64% from widespread lowland Neotropical sources; 17% from western Mexico and central America, 4% from Venezuela, Colombia, or northern Ecuador; 4% known only from Ecuador; 6% from semi-arid coastal Ecuador, Peru, Chile or arid Argentina; 1 % West Indies; 4% from western Pacific (Austral-Asian) region. In plants, an Andean highland component is present but this is not evident in insects. The transport of the ancestors of the endemics was mostly through the air as winged adults. Fewer (especially the wingless species) arrived by sea, rafting on floating materials. The smallest component arrived as ecto- or endoparasites on insect or vertebrate hosts (Peck 1996).

The distributions of the endemic insect species are similar to the patterns in the plants. In insects, the endemics are usually restricted to a single island, and this is more prevalent for humid-zone species. The endemics on more than one island probably represent natural dispersal of the species after its origin on a single island. Although most endemic invertebrates are single-island species, on average, endemic species tend to be found on more islands than non-endemic native species, even more of which are restricted to a single island (Peck 1996).


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Summary

• The biodiversity (abundance and distribution of species) of Galapagos is the least altered of any oceanic archipelago.

• Galapagos demonstrates conspicuous and abundant examples of evolutionary processes.

• The flora and fauna of Galapagos are globally renowned — such as Darwin’s finches, giant tortoises, relatives of daisies that have evolved into trees, marine iguanas etc.

• Galapagos is an area of high endemism for many taxonomic groups, including snails, trees and shrubs, insects and vertebrates.

• The islands and their biodiversity represent an international and historical monument of intellectual development and cultural change.

REFERENCES

Baert, L. 2000. Invertebrate research overview: 1. Terrestrial arthropods. Pp. 23-25. in Science for conservation in Galapagos. Ed. N. Sitwell. Bulletin de L’Institut Royal des Sciences Naturelles de Belgique, Vol. 70-Supplement.

Chambers, S.M. 1991. Biogeography of land snails. Pp. 307-326. In Galapagos Invertebrates: Taxonomy, biogeography and evolution in Darwin’s islands. Ed. M.J. James. Plenum Press, N.Y.

Colinvaux, P.A. 1984. The Galápagos climate: present and past. Pp. 55-69 in Key Environments. Galápagos. Ed. by R. Perry. Pergamon Press, Oxford.

Crawford, D.J., Whitkus, R. & Stuessy, T.F. 1987. Plant evolution and speciation on oceanic islands. Pp. 183–199 in Urbanska, K.M. (ed.) Differentiation patterns in higher plants. Academic Press, New York.

Elisens, W.J. 1989. Genetic variation and evolution of the Galapagos shrub snapdragon. Nat. Geogr. Res. 5: 98–110.

Finston, T. & Peck, S.B. 1995. Population structure and gene flow in Stomion; a species swarm of flightless beetles of the Galapagos beetles. Heredity 75(4): 390-397.

Geist, D. 1996. On the emergence and submergence of the Galápagos islands. Noticias de Galápagos 56: 5–9.

Lawesson, J.E., Adsersen, H. & Bentley, P. 1987. An updated and annotated checklist of the vascular plants of the Galapagos islands. Reports from the Botanical Institute, University of Aarhus 16.

Lowrey, T.K. & Crawford, D.J. 1985. Allozyme divergence and evolution in Tetramolopium (Compositae-Astereae) on the Hawaiian islands. Syst. Bot. 10: 64–72.

Peck, S.B. 1996. Origin and development of an insect fauna on a remote archipelago: The Galapagos Islands, Ecuador. Pp. 91-122. In The Origin and evolution of Pacific island biotas, New Guinea to Eastern Polynesia: patterns and processes. Ed. by A. Keast and S.E. Miller. Academic Publishing, Amsterdam.

Peck, S.B. 1997. The species-scape of Galapagos organisms. Noticias de Galapagos 58: 18-21


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Peck, S.B., Heraty, J., Landry, B., and B.J. Sinclair. 1998. Introduced insect fauna of an oceanic archipelago: the Galapagos Islands, Ecuador. American Entomologist Winter Issue: 219-237.

Porter, D.M. 1983. Vascular plants of the Galapagos: origins and dispersal. In Patterns of Evolution in Galapagos Organisms (eds R.I. Bowman, M. Berson & A.E. Leviton) pp. 33–96. American Association for the Advancement of Science, San Francisco.

Rick, C.M. & Fobes, J.F. 1975. Allozymes of Galápagos tomatoes: polymorphism, geographic distribution, and affinities. Evolution 29: 443–457.

Schatz, H. 1991. Catalogue of known species of Acari from the Galapagos Islands (Ecuador, Pacific Ocean). Int. J. Acarology 17: 213–225.

Snell, H. M., P. A. Stone, and H. L. Snell. 1996. A summary of geographic characteristics of the Galapagos Islands. Journal of Biogeography 23: 619-624.

Snell, H.L., and S. Rea. 1999. El Niño 1997 – 1998 en Galápagos: ¿Se puede estimar 120 años de variaciones climáticos con estadísticas de 34? En: P. Ospina y E. Muñoz (eds) Informe Galápagos 1998 – 1999, pp 65-71. Fundación Natura, Quito, Ecuador.

Weber, W.A. & Gradstein, S.R. 1984. Lichens and Bryophytes. pp. 71-84 in Key Environments. Galápagos. Ed. by R. Perry. Pergamon Press.

CDF/WWF Biodiversity Vision Chapter 4 – Terrestrial Conservation Criteria

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CHAPTER 4 – CONSERVATION CRITERIA FOR THE TERRESTRIAL BIOME Principal Authors – A. Tye and H.L. Snell

Galapagos is unusual as an oceanic archipelago in that we can still accurately identify the pre-human biological status. Very few taxa have been lost, and it would be theoretically possible to restore Galapagos to very nearly complete pre-discovery conditions. If the biodiversity vision is to maintain and restore as much as possible of the native biodiversity and natural ecological and evolutionary processes, then we need to establish measures of our success or failure in achieving this goal (Chapter 6).

4.1 General Criteria

Measuring the success of conservation may be approached in many ways. Here we combine three levels of biological scale (species, communities/habitats, and landscape) with two biological processes (ecological and evolutionary) and an important condition (the status of alien species) to develop a set of criteria that will measure progress towards the conservation of biological diversity in the Galápagos archipelago (adapted from Noss 1991). The fact that “ecosystems” are omitted is not meant to belittle their importance, rather it stems from the observation that precise measures of ecosystem metrics are extremely difficult in the Galapagos. In general it seems that most of what was proposed by workshop participants more closely reflected measures of habitat rather than ecosystem. Periodic evaluation of these criteria will establish the status and likely trajectories of change for the foreseeable future (see Chapter 6). Questions within these criteria are:

4.1.1 Species. To what extent are viable populations of all native species maintained in natural patterns of abundance, distribution and variation? This “species-level” criterion should be understood to include variation within species (infraspecific taxa, distinct populations, and the full range of genetic and phenotypic variation of each taxon).

4.1.2 Communities/Habitats. To what extent are all distinct natural communities, habitat types and seral stages represented, across their natural ranges of variation, in protected areas insulated from anthropogenic impacts, including the introduction of alien species?

4.1.3 Landscape/Habitat Extent. To what extent are blocks of natural habitat, large enough to be resilient to large-scale periodic natural disturbances and long-term changes, maintained free from significant impacts of anthropogenic factors?

4.1.4 Processes. To what extent are ecological and evolutionary processes that create and sustain biodiversity, including biotic interactions, nutrient cycles, natural disturbance regimes, patterns of geneflow, dispersal, and microevolutionary change maintained?

4.1.5 Alien species. How successful is the prevention of anthropogenic introduction and establishment of alien species; and the control and eradication of established aliens or the mitigation of their effects on native biodiversity?

4.2 Specific Measures

For each of the above general criteria, several different measures should be evaluated. The following list presents selected variables that can be measured for each of these five criteria, in order to answer the above questions. The list is not exhaustive and other variables may be included. In general “baseline” values refer to those assumed in 1535. They may reflect the 1999 or some other reference condition believed to be either natural (pre-discovery) or desirable. A subset drawn from these specific measures forms the framework of projections for the future status of terrestrial biodiversity in Chapter 6.

4.2.1 Species. Specific measures within this criterion require some general explanation. Throughout this document the authors have tried to standardize use of some confusing terminology. We use the term “native” to denote natural occurrence of components of biodiversity. In this use “native” includes endemic and non-endemic but naturally occurring


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forms. We also refer to species, infraspecific taxa and distinct populations. The three terms are necessary to encompass the range of variability potentially managed in the conservation biology of the Galapagos. “Species” refers to formally named species. “Infraspecific taxa” refers to named subspecies, races, or varieties. “Distinct populations” refers to apparently evolutionarily distinct populations that are not taxonomically recognized. Within different groups of organisms the degree to which these terms are used varies. Some of these measures are loosely adapted from IUCN (1994).

• Numbers of native species, infraspecific taxa, or distinct populations extant/extinct.

• Numbers of native species, infraspecific taxa, or distinct populations assessed as “Threatened” (see IUCN categories reproduced in Annex 4.1).

• Abundance of each native species, infraspecific taxon, or distinct population (with reference to baseline abundance). In general, abundance will often be monitored by changes in relative density.

• Metapopulation structure of each native species, infraspecific taxa, or distinct population (with reference to baseline structure). Metapopulation structure includes the number of subpopulations, demes, and movements among them.

• Area and distribution of occurrence for each native species, infraspecific taxon, or distinct population (with reference to baseline area and distribution). This measure encompasses the spatial component of biodiversity. The size of the area occupied by a form has obvious relevance to its status. However, where that area is located is also significant. If a species has been extirpated from its natural range but introduced to another area (Baltra land iguanas are a good example) its status is different than a species remaining within its natural range.

• Population structure (age, size structure) of key native species, infraspecific taxa, or distinct populations (with reference to baseline or stable structures). This measure provides a possible “early warning signal” of future decline. It is more difficult to measure than those listed previously and therefore is limited to key forms.

• Genetic and phenotypic variability of selected indicator native species (with reference to baseline level of variation). Changes in genetic and phenotypic variability can provide an “early warning signal” of longer processes leading to declines in biodiversity. As with population structure it is a difficult measure to assess and will therefore be limited to a subset of species.

4.2.2 Communities/Ecosystems. Within specific measures of this criterion “communities” refers to populations of individuals of species with the potential of typical biotic interactions (predation, competition, herbivory, parasitism, mutualism, etc.). “Ecosystems” include the communities and the abiotic environmental processes that affect life. Physical processes included with general areas such as hydrology, geology, oceanology, and climatology are implicitly included.

• Number of natural communities represented in areas insulated from anthropogenic impacts.

• Area and distribution of each natural community. See discussion of “area” and “distribution” under 4.2.1.

• Changes in species composition of communities attributable to anthropogenic factors.

4.2.3 Landscape/Habitat Extent. Specific measures of this criterion attempt to quantify changes at the broadest scale of biodiversity. They have a significant spatial component and are the measures that most frequently address “what is where.” They are adapted from Bennett (1999).

• Total spatial extent (distribution & area) of each key habitat type (measures habitat loss).


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• Fragmentation of blocks of key habitat types (measures habitat and population fragmentation).

• Reduction in size of blocks of key habitat types (measures habitat reduction and size of subpopulations).

• Increased isolation of habitat fragments from one another (measures habitat isolation: distance between subpopulations).

• Numbers and sizes of “pristine” islands. At the landscape scale much of the variation within the specific measures mentioned above can be encompassed in the number of “pristine” islands. A pristine island would be one with all of its native biodiversity and no alien species. However, there are several species of widely dispersed plants and invertebrates that appear to be minor threats to the natural biodiversity of Galapagos. We distinguish between “pristine”, “near pristine” and “not pristine” islands. Islands with no species of alien vertebrate, fewer than 5 species of invasive alien plants, where there are no unnatural extirpations and no large-scale anthropogenic changes to the ecosystem, are considered “near pristine”.

4.2.4 Ecological and evolutionary processes. While there were many ecological and evolutionary processes discussed in the workshop, we quickly found that the specific measures discussed concentrated on anthropogenic activities predicted to impact those processes rather than the processes themselves. The following specific measures are a subset of process components that can be directly measured.

• The relative occurrence of altered biotic interactions (compared to the baseline). Altered biotic interactions can involve alien species in significant ecological roles (alien predators, competitors, herbivores, parasites, or mutualists for example). Altered biotic interactions would also include native species with unnatural ecological roles, or situations where natural roles gain unnatural significance. Natural predation by Galapagos hawks on young tortoises is a potential example. It appears that hawks have always eaten young tortoises, but the rate of predation is naturally insignificant. However, when a tortoise population declines sharply to a very low level, the rate of predation by hawks becomes a primary force preventing recruitment of juveniles into the adult population.

• The occurrence of unnatural disturbances or anthropogenic climate change. In general we are concentrating on measures that quantify impacts with some level of control within Ecuador or the Galapagos. Thus we are not proposing measures of impacts of global climatic change. Smaller scale unnatural disturbances like fire or fuel spills are the specific measures of unnatural disturbance we propose. Microclimatic change as a response to environmental modification are certainly of concern.

• The occurrence of microevolutionary change caused by unnatural selective forces, altered rates of geneflow, heightened mutation rates, or artificial bottlenecks. The specific measures most likely to be used are those concerning unnatural selective forces and altered rates of geneflow. Those two are often caused by reduced isolation or increased habitat fragmentation and artificial selective agents like alien species. Measures of the effects of artificial bottlenecks have already been covered in 4.2.1 under genetic variation.

4.2.5 Alien species. The extreme importance of alien species as threats to the biodiversity of the Galapagos makes specific measures directly related to them potentially helpful in gauging the success of conservation efforts. There was considerable discussion in the workshop about using measures of human activity that correlated with aspects of the problem of alien species, similar to the discussions about correlates rather than processes in section 4.2.4. Here the authors have concentrated on the direct measures and refer readers to the table of “driving forces” in Chapter 6 for a subset of measures of the correlates.

• Rates of arrival and establishment of alien species (including pathogens).

• Abundance and distribution of alien species.


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• Number of alien species eradicated. This serves to quantify restoration efforts.

• Number of alien species whose distribution and/or abundance have been reduced (number of species, amount of reduction). This quantifies the control effort for those species where eradication seems unlikely.

• Change in abundance or distribution of endemic/native species directly related to the effects of alien species.

• Success of specific mitigation efforts for native species impacted by aliens that can’t be eradicated. This is an interaction between alien species and native species criteria. Certain types of mitigation efforts may not require altering sizes of alien populations. “Headstarting” of Galapagos tortoises is a good example. The assumed effects of alien black rats on Pinzon Island (apparent depredation of all hatchling tortoises) are mitigated by raising hatchling tortoises in captivity until they are large enough to avoid predation by black rats. Here the criteria of success is a tortoise population with a positive growth rate even though the numbers of rats remains high.

4.3 Specific Components of Diversity to Serve as Indicators

Within each of the specific measures (4.2) proposed for the five general criteria (4.1), it will not be possible to monitor every species, habitat or site and it will be necessary to choose representative indicators. In some cases, a variable to be measured can be more specifically directed at an island, habitat, area or species. The following is a general presentation of possibilities. One of the important activities for the immediate future is a more rigorous evaluation of potential indicators.

4.3.1 Species. Key indicator species might include groups with large scale radiations such as the plant families Asteraceae, Cactaceae and Amaranthaceae, the Bulimulidae land snails, terrestrial Isopoda, vertebrates such as the tortoises, finches, lava lizards and rodents. Indicator species should also include species or groups sensitive to a range of environmental impacts, such as phytophagous insects or birds. They should include a wide range of taxonomic diversity and trophic levels, as well as species present in key sites or key habitats, such as snakes, bats, predatory birds, seabirds, marine iguanas and soil microfauna and flora. Species facing local extinctions or genetic introgression with introduced species should be included, such as the Galapagos tomato Solanum cheesmaniae, as well as those subject to direct exploitation, such as timber trees. We anticipate that introduced pathogens and disease will play increasingly important roles, so species such as birds with known sensitivities to these potential new threats will be important. Where possible, indicator species and groups would be chosen for ease of monitoring.

4.3.2 Communities/Habitats. Communities to serve as indicators should include those under special threat, including especially the humid highlands. Monitoring should include representative habitat types of broader zones subject to a wide range of environmental impacts. Listed by zone, these include:

• Littoral: mangroves, brackish lagoons, beach ridges, sand dunes and salt marsh;

• Arid: scrub communities dominated by Cordia, Prosopis, Bursera, Gossypium and Croton, Opuntia/Scalesia on lava outcrops;

• Transition: woodland communities dominated by Pisonia, Psidium, Psidium/Scalesia;

• Scalesia: communities with understory dominated by Psychotria, Alternanthera halimifolia;

• Brown: dominated by Frullania;

• Miconia (and other wet highland scrub): communities dominated by Miconia, Acnistus , Croton;


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• Fern-sedge: communities dominated by grasses and sedges (Ciperacea), Pteridium, groves of tree-ferns;

• Dry highlands: dry highland scrub, dry highland grassland;

• Freshwater habitats: bogs, vertical bogs, marshes, wet meadows, streams, waterholes, lakes, temporary ponds;

• Pioneer series: Brachycereus, Jasminocereus/Darwiniothamnus/Scalesia, ash plains, fumaroles, Macraea scrub.

4.3.3 Landscape/Habitat Extent. Although indicators for the Landscape/habitat extent monitoring should focus on the most threatened islands, especially the inhabited or formerly inhabited ones, islands should be chosen to be representative of a variety of levels of biodiversity conservation status, from near-pristine to heavily damaged, and a variety of specific pressures, such as grazing by introduced herbivores. Selected islands might include: Fernandina, Pinta, Marchena, all inhabited islands (and northern Isabela), Santiago. Remote sensing could be the key technique for monitoring at the landscape level. A specific research requirement is preparation of good vegetation maps for the entire archipelago.

4.3.4. Processes. Situations that could provide insight into changes of the ecological process criterion could include herbivory by tortoises and alien species; predation by snakes, hawks, and aliens such as cats, dogs, and pigs; parasitism by alien invertebrates and mutualisms between tortoises and alien plants (tortoises disperse seeds of alien plants for example). Specific indicators for the evolutionary processes criterion should include those with known examples (i.e. body size and behavior in lizards) and significant potential (artificial bottlenecks in population size for Opuntia, tortoises, iguanas, and hawks). Altered patterns of geneflow as a response to altered isolation could be examined in native species know to be commonly moved among islands (geckos, arachnids, grasses, etc).

4.3.5. Alien species. Potential indicators within the alien species criterion should focus on three areas: species already introduced, potential new arrivals, and mitigation of impacts for those where eradication is unlikely or postponed. For new arrivals, lists of unwanted species should be prepared, but all species are regarded as prohibited unless specifically evaluated as safe for introduction. Monitoring should also concentrate on the most dangerous (invasive, aggressive) organisms. Specific focus groups include aquatic plants, vines, trees, bushes, grasses, ants, phytophagous and predatory insects, terrestrial and freshwater molluscs, other freshwater invertebrates, snakes, mammals, pathogens of plants and animals. Focal sites include ports of entry, agricultural zones, towns. Key sites for monitoring alien species largely coincide with key sites for monitoring at the species, community and landscape levels, such as near-pristine islands, sites with concentrations of endemic species etc. Other monitoring programmes will produce data at more than one level, such as interactions between native and introduced species, including species composition changes. Alien species that affect critical native species should also be selected for monitoring.

4.4 Research needs & summary

Some of the above variables are far easier to measure than others and it is not anticipated that values of all variables will be available for assessment at each future evaluation. However, an attempt should be made to establish research and monitoring programs that permit assessment of as many of the above variables as possible and are designed with these questions in mind.

A crucial deficiency is that for many of the above variables we lack current baseline data. An urgent research priority must therefore be to establish baseline values for as many of these variables as possible.


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REFERENCES

Bennett, A.F. (1999) Linkages in the Landscape. IUCN, Gland.

IUCN (1994) IUCN Red List Categories. IUCN, Gland.

Noss, R. F. (1991) Protecting Habitats and Biodiversity, Part 1: Guidelines for Regional Reserve Systems. National Audubon Society, New York.

CDF/WWF Biodiversity Vision Chapter 5 – Status of Terrestrial Biodiversity

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CHAPTER 5 - CURRENT STATUS OF AND THREATS TO THE TERRESTRIAL BIODIVERSITY OF GALAPAGOS Principal Authors – H.L. Snell, A. Tye, C.E. Causton and R. Bensted-Smith

In this chapter the authors discuss the status of Galapagos terrestrial biodiversity at the time of the workshop (1999) with updates to 2001 where appropriate. We discuss the changes that have taken place since 1535, and the anthropogenic factors that have brought about those changes.

STATUS OF GALAPAGOS BIOLOGICAL DIVERSITY IN A GLOBAL CONTEXT

“The Galápagos Islands are unique among oceanic archipelagos in that over 95 % of their biological diversity remains recoverable” is a common sentiment within the conservation community. The statement is true, but only because of the relatively short duration of human presence in the Islands. Many other oceanic archipelagos have been inhabited for thousands of years. The majority experienced strong waves of “prehistoric” anthropogenic extinctions that are not recorded. Because these early extinctions occurred prior to recorded history there is no baseline to measure against. The magnitude of these extinctions can only be estimated from subfossil remains and archeological analysis. The Galapagos escaped these extinctions simply because they were not colonized by humans. Unfortunately, once colonized, the Galapagos seem to be following a similar trajectory to that followed long ago by other archipelagos.

It is often assumed that the current rate of extinctions in the Galápagos is low compared to other areas of the world, but this is not the case. If we compare extinctions within the period of recorded biological history (about the last 400 years), few countries or archipelagos have rates of extinction higher than Galápagos. Part of that pattern can be accounted for by the general tendency of islands to have more historic extinctions than continental areas, but the Galápagos rank high even among other islands. When compared by area, nearly 90% of a sample of 95 oceanic islands have had relatively fewer extinctions per km2 than Galápagos in the past 400 years (Figure 5.1). When compared by biological diversity, approximately 81 % of the same sample of oceanic islands have lost a lower proportion of their species diversity in the past 400 years (Figure 5.1). It is important to note that the sample of oceanic islands does not include every archipelago in the world (it was drawn from Groombridge 1992 and Groombridge & Jenkins 1994).

Historic Extinction vsArea

Area (km2)

10-1 100 101 102 103 104 105 106 107 108

Ext

inct

ions

: Ver

tebr

ates

& P

lant

s

100

101

102

ContinentalOceanic IslandsGalápagos

ContinentalOceanic IslandsGalápagos

Historic Extinction vsDiversity

Species Richness: Vertebrates & Plants101 102 103 104 105

Ext

inct

ions

: Ver

tebr

ates

& P

lant

s

100

101

102

Figure 5.1. Global Patterns of Extinction Within the Last 400 Years. Numbers of recorded extinctions per geopolitical unit for mammals, birds, reptiles and plants combined. See Chapter 2 for explanation of data.


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The general message from this analysis is that the Galapagos do represent a unique opportunity to preserve an oceanic archipelago’s biological diversity, but it is an opportunity that will be quickly lost without prompt action. The remaining sections of this chapter examine the extinctions, population declines, and threats that must be solved to realize the opportunity.

TERRESTRIAL STATUS AND THREATS

As explained in Chapter 4, criteria may be chosen to measure change at several different levels. Owing to limitations of available baseline data and the difficulty of measuring certain criteria, especially at the landscape and ecological-evolutionary process levels, the discussion below deals primarily with measures at the species-population, community-habitat and alien species levels. Although the authors recognize the importance of other measures, we are unable at present to quantify them extensively enough for use within this publication. Within the Galapagos, our basic geographic units are islands. Where possible we treat major habitat types within islands, but in many cases the data were limited to island-level accuracy. It is important to recognize the difference between the data available to establish losses and changes between 1535 and 1999 and that necessary to monitor and manage biological diversity in the future. The fact that most of the following is limited in scope and often appears to overlook criteria set out in Chapter Four, is not an oversight. Hopefully the next time the biological diversity of Galapagos is reviewed, data for all levels of criteria will be available.

GALAPAGOS IN 1999

Extirpations (species, populations)

Galapagos presents a typical rate of natural extinction, due to natural processes such as (non-anthropogenic) climate change and volcanic eruptions, but contemporary extinction in all groups of plants and animals has mostly been caused by humans.

Galapagos has suffered rates of recent anthropogenic extinction comparable to other oceanic archipelagos, but its prehistoric rate of anthropogenic extinction is zero, because of its relatively recent discovery and settlement by man. Three well-documented endemic species of vascular plant have become extinct (Blutaparon rigidum on Santiago and Sicyos villosa and Delilia inelegans on Floreana). The losses of entire plant species have been few, and Galapagos has similarly lost few infraspecific populations of plants from individual islands or sites. However, many more individual populations, as well as entire species, have been brought to the edge of extinction by human-mediated change (see below: Threatened Species).

There is no firm evidence of extinctions among the terrestrial invertebrate fauna, other than for the endemic family of land snails, the Bulimulidae, of which some have undoubtedly gone extinct although we do not know how many. For example, only one live Bulimulus achaetellinus has been collected since 1890 (in 1987) (Coppois 2000). Other invertebrate species which may now be extinct include the Neoryctes scarab beetles (Cook et al. 1995). There are undoubtedly other extirpations among invertebrates that have gone unrecorded, because baseline data for invertebrates have, for the most part, only become available following intensive surveys in the last 30 years (Baert 2000, Peck 1996). Undoubtedly some other invertebrate extirpations or extinctions have occurred, but they presently seem to be few and poorly documented, and the evidence is usually indirect. For example, small endemic terrestrial isopods have not been found recently on the large islands where they were formerly recorded, with all recent records coming only from small islets that have remained free from introduced species of terrestrial isopods. Local populations of some invertebrates such as spiders, scorpions and native ant species can no longer be found in areas infested with introduced Wasmannia auropunctata ants (Lubin 1984, Roque et al. 2000). Opuntia-feeding Gerstaeckeria weevils have not recently been found on some of the islands where their host plants have virtually vanished due to browsing pressure from feral donkeys and other herbivores. Other host-specific phytophagous insects may have experienced similar population declines or extinction. The seeming rarity of large flightless scarab beetles could be due to predation on them by introduced mice and rats.

The vertebrates of Galapagos have by far the greatest number of recorded extirpations, approximately nine percent of the baseline vertebrate fauna appears extinct. Of the oceanic archipelagos analyzed (see chapter 3), only four have had a greater number of recent extinctions of vertebrates, (counting Hawaii, not in the sample, there would be five). This


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pattern could partially result from the greater attention paid to vertebrates, but it also appears that several components of their life-history may make them more susceptible than many plants and invertebrates.

The ten recently extinct vertebrates of the Galapagos include two tortoises (Geochelone phantastica of Fernandina, and Geochelone elephantopus of Floreana), two giant rice rats (Megaoryzomys curioi of Santa Cruz and Megaoryzomys sp of Isabela), five rice rats (Oryzomys galapagoensis of San Cristobal, two Nesoryzomys of Isabela, and Nesoryzomys darwini and Nesoryzomys indefessus of Santa Cruz), and a land iguana (Conolophus sp of Santiago). There are two other tortoises (from Rabida and Santa Fe) occasionally listed as extinct. However, the Rabida tortoise was not a distinct form and the tortoises of Santa Fe were probably introduced. In addition to these “species-level extinctions” there are many extirpations of island populations including snakes, hawks, several terrestrial birds, and possibly a gecko.

The vertebrate extirpations are the result of over-exploitation and interactions with introduced species. Galapagos tortoises and hawks are the most susceptible to overexploitation. Hypotheses about the extinctions of rice rats often involve diseases carried by black rats or direct competition, but recent data suggest that feral cats may have played a greater role than previously appreciated. Potential causes for the local extirpations of several terrestrial birds include predation (feral cats and rats) and introduced diseases.

Threatened species

A review of the threat status of the endemic flora, using IUCN categories and criteria (described in Annex 4.1), was completed in 2001, with all 175 endemic species having been assessed, of which 8 could not be adequately treated (Data Deficient) (Tye 2002). Table 5.1 demonstrates that, of the taxa fully evaluated, almost 10% are classed as Critically Endangered, i.e. they are already reduced to the brink of extinction. Some 15% are Endangered, meaning that they are in serious decline, and a further 40% are classed as Vulnerable, in most cases due to their very small ranges. Compared with the small number of plant extinctions so far, the salient point from these figures is that a relatively large number of species (particularly those in the categories CR and EN, and some of the VU) have suffered decline, are continuing to decline, and are thus on the road to extinction unless action be taken to reduce or remove the threat factors leading to those declines. Even more species (the remainder of those classed as VU) are extremely susceptible to rapid declines and loss caused by new threat factors, such as introduced pests and diseases.

Table 5.1. Numbers and percentages (in parentheses) of vascular plant taxa in each IUCN threat category (from Tye 2002).

Threatened

Taxa fully evaluated

EX CR EN VU NT LC

Species 167 3 (2%) 13 (8%) 21 (13%) 61 (37%) 15 (9%) 54 (32%)

All taxa1 220 3 (1%) 19 (9%) 32 (15%) 87 (40%) 16 (7%) 63 (29%)

EX=Extinct; CR= Critically Endangered; EN=Endangered; VU=Vulnerable; NT=Near-threatened; LC=Least Concern. No taxa were classified as EW (Extinct in the Wild).1Excludes species with infraspecific taxa in order to avoid inflating numbers artificially.

The figures for current status given in Table 5.1 compare with a situation in 1535 where no species would have been classed as CR or EN, although the number classed as VU would have been non-zero and perhaps comparable with the present figure (including species with small ranges, and probably all those now classed as EX, CR and EN).

No formal assessment has been made of decline in individual populations of plants, where they have not been named as taxa. However, many populations are known to have declined drastically, especially as a result of damage caused by introduced mammals and by clearance of their sites for agriculture. Examples include several species that are no longer present, or are


33

severely reduced, on islands which have or have had large populations of introduced mammals (e.g. Lecocarpus lecocarpoides on Española), whereas the (naturally smaller) populations on their neighboring islets remain at high density. Scalesia pedunculata is almost extinct on San Cristóbal, because of the conversion of almost its entire range to agriculture, and it is now rare on Santiago because of damage by goats, although it remains fairly common (though much reduced) on Santa Cruz and Floreana. The genetic distinctness, and evolutionary value, of the severely reduced populations in such cases is rarely known.

Since little is known about population numbers and distribution of most terrestrial invertebrate species, it is hard to identify threatened species. Limited and circumstantial evidence leads us to believe that many would qualify as Threatened by IUCN criteria. There are many accounts of declines reported and up to 60% of all arthropod species may have suffered declines, with the greatest losses in the Humid Zone (see below). In the “small orders” of insects, representing some 484 species in total, 140 species occur in the humid forest zone, of which 51 are archipelago endemics, and 25 are single island endemics. This zone has experienced the greatest alteration caused by humans, most taking place before recent insect diversity surveys. The alteration of this habitat zone has undoubtedly influenced insect species diversity and abundance. Species that are endangered or are most likely to become endangered include the phytophagous endemics that have become specialized on threatened plant taxa, such as Lepidoptera that feed on Scalesia, and the sphingid moth Xilophanes tersi, which feeds on Psychotria. Ectoparasites of endemic vertebrates, such as those of the rice rats, may also have declined. Many invertebrate species are under threat in the smaller islands where the fire ants Wasmannia auropunctata and Solenopsis geminata have been introduced, such as Champion and Albany islets.

Fifty percent of the surviving vertebrate fauna of Galápagos is likely to go extinct if current and future conservation efforts are not successful (Table 5.2). Birds appear to have potential for the greatest increase in extinction, largely because the most vulnerable of the other vertebrates are already extinct. Interactions with alien species will increase in importance as causes of extirpations, but overexploitation remains the greatest threat for one species of Galapagos tortoise and Galapagos hawks.

Table 5.2. Numbers and percentages (in parentheses) of vertebrate taxa in IUCN threat categories.

Threatened

Total taxa evaluated

EX EW CR EN VU NT LC

Species 112 10 (9) 1 (1) 4 (4) 12 (11) 38 (34) 5 (4) 42 (38)

EX=Extinct; EW=Extinct in the Wild, CR= Critically Endangered; EN=Endangered; VU=Vulnerable; NT=Lower Risk (near-threatened); LC=Lower Risk (least concern).

THE THREATS

In this section we discuss the major human activities that threaten the persistence of the natural biological diversity of the Galapagos. In the terrestrial arena these are habitat degradation or alteration, overexploitation ( i.e. non sustainable extractive use of natural resources) and interactions with alien species. The fact is often quoted that some 96% of Galapagos’s land area is included within the Galapagos National Park, which gives the impression that terrestrial habitats are well protected. In this section, we demonstrate that this figure is misleading, and that the impact of the small areas excluded from the national park is out of all proportion to their comparatively tiny size.

Habitat Alterations

Of the four broad vegetation zones discussed in Chapter 3, the Littoral and Humid Zones are the most restricted, but the Littoral Zone contains little biological diversity compared to the


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highly diverse Humid Zone with its many endemic species. The Littoral Zone occurs on most islands, irrespective of their size, whereas the Humid Zone occurs only on the larger, higher islands, and has been attractive to human settlement, as it has more available water and is the most suitable zone for agriculture.

Islands with Humid Zone Inhabited? Floreana Yes Isabela Yes San Cristóbal Yes Santa Cruz Yes Santiago Formerly Fernandina No Pinta No

Most islands with a humid zone are or have been inhabited.

The major part of the 4% of Galapagos land area that is settled is in the highlands. The major human activity in the higher parts of the islands is agriculture. On each of the islands with a civilian population there is an Agricultural Zone, all of which are in the highlands.

Map of Floreana Island showing the major vegetation zones and the Agricultural Zone.


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Map of Santa Cruz Island showing the major vegetation zones and the Agricultural Zone.

Map of southern Isabela Island showing the major vegetation zones and the Agricultural Zone.


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Map of San Cristóbal Island showing the major vegetation zones and the Agricultural Zone.

Habitat clearance has thus been most damaging to the Humid Zone, as shown by the data in Table 5.3. In particular, San Cristobal has lost almost all of its Humid Zone, and Santa Cruz almost 75%. On Floreana, the proportion of the Humid Zone cleared for agriculture is small, but the total area of Humid Zone on that island is also small, and other impacts of agriculture (principally introduced species) have severely affected the remainder. Indeed, the remainder of the Humid Zone not occupied by agriculture, on all inhabited islands, has been more severely damaged and is facing the greatest threat from introduced species escaping from the Agricultural Zones, than any other vegetation zone. Further, we must remember that the Humid Zone includes several quite distinct sub-zones that, in many cases, have been much more severely reduced than the Humid Zone as a whole. For example, San Cristobal and Santa Cruz have lost nearly 100% of their former Scalesia Zones. Finally, on every island there are single-island endemic species, especially of plants and invertebrates, restricted to one single Humid Zone of one island; many of these, especially on the worst-affected islands, must now be on the edge of extinction.

The conclusion must be that it is seriously misleading to speak of 96% of Galapagos land area being protected, without qualification.


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Table 5.3. Habitat clearance for agriculture, and habitat degradation by introduced plants and herbivores, on the four inhabited islands. Status in 1999.

Island Zone

Total area of habitat type (km2)

Area in km2 and (%) of this habitat occupied by Ag. Zone

Area and (%) of this habitat uncleared

Estimated % uncleared and with species composition not significantly altered

Santa Cruz1 humid 118 87 (74) 31 (26) < 5

transition 127 33 (26) 94 (74) 30–50

agricultural 122 [100]

Floreana2 humid 31 5 (15) 26 (85) 10–20

transition 39 1 (2) 38 (98) 20–30


San Cristóbal humid 83 77 (93) 6 (7) < 1

transition 40 4 (9) 36 (91) 30–50


Isabela1,3 humid 641 52 (8) 589 (92) 30–40

transition 1323 0 (0) 1323 (100) 30–40


Sierra Negra1,3 humid 370 52 (14) 318(86) 20–30

transition 460 0 (0) 460(100) 20–40


Santiago humid 35 < 1 (1) (Park huts) 35 (99) 5–10

transition 32 0 (0) 32 (100) < 10

Pinta humid 4 0 (0) 4 (100) 5–10

transition 6 0 (0) 6 (100) < 10

Fernandina humid 105 0 (0) 105 (100) 100

transition 143 0 (0) 143 (100) 100

1 Note that the proportions of the humid zone affected by agriculture on Santa Cruz, Isabela (including Sierra Negra) are greater than shown by these figures, due to the former presence of farms (of undetermined extent) in areas outside the present Agricultural Zones; further, on Isabela (including Sierra Negra) large areas have been invaded and dominated by guava Psidium guajava, outside the Agricultural Zones.

2 The figures for Floreana are the least accurate in this table, owing to inadequacies in existing maps. The agricultural zone is probably larger than shown, and the overlap with the humid zone probably greater.

3 Sierra Negra is the volcano on Isabela where the human settlements are located.


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Extractive Use

The inhabitants of Galapagos have turned to its natural resources for many of their needs. In the terrestrial realm, such direct exploitation has affected relatively few species, but the effects on those that have been used have often been severe. Few native plants are of direct use to humans, but trees have been widely used for their timber and in early days for firewood. Species that have been favoured have often become either very rare (e.g. Lippia salicifolia, endemic to Floreana), or their population structure has been changed in such a way as to cause possible and as yet unknown effects on their evolution. Perhaps the most sought-after Galapagos timber is Matazarno Piscidia carthagenensis, valued for its strength and resistance to rot and insect attack. On all islands where it occurs, most of the large specimens, with the best form, have already been harvested. This may have several effects. Removal of the “best” specimens, takes out a selected sample of genetic variation, changing the genetic make-up of the species. Given that the species is extremely slow growing, removal of so many adults will affect the population structure far into the future, with unknown long-term effects on recruitment and the equilibrium of the species with its natural predators (such as seed-eating beetles). Since the decline in Matazarno, loggers have turned to the endemic Guayabillo Psidium galapageium, which may soon lead to similar effects on this species.

Extractive use of mineral resources can have severe effects on certain species. Quarries for road-stone and building usually exploit scoria cones. These have been shown to harbour unique populations of Bulimulid snails, with entire species restricted to a single cone (Coppois 1984, Coppois & Wells 1987). Given the dearth of distributional studies of invertebrates, the extent to which such species have been endangered or even extinguished by such activities is unknown. In a few cases, mineral extraction has also endangered plants. The Critically Endangered Linum cratericola was only discovered in 1966, and has only ever been known from two sites, both on Floreana. One of these was severely damaged by a road-stone quarry, and the species is now extinct there, probably at least partly due to habitat loss. Calandrinia galapagosa, another Critically Endangered plant endemic to San Cristóbal, has its major remaining site on the outer wall of a scoria cone that was, until 1999, being excavated from the other side by a road-stone quarry. This cone is also a major site for another threatened Cristóbal endemic, Lecocarpus darwinii. There are plans to open a new quarry at another important site for this latter species.

By far the most serious example of unsustainable exploitation of terrestrial Galapagos organisms comes from the tortoises. Prior to 1900 the majority of human terrestrial activity in the islands involved killing tortoises. Reasonable estimates for the numbers of tortoises removed range from several hundred thousand to a million tortoises. All populations of tortoises on all islands were exploited, most were reduced by more than 60 percent, two were effectively extirpated, and none have fully recovered 100 years later. Unfortunately, unsustainable exploitation of tortoises remains the greatest threat to the survival of two species on Southern Isabela (Geochelone vicina and G. guntheri). In the rest of the archipelago efforts to prevent human predation of tortoises are effective.

A variety of other terrestrial vertebrates have been exploited at unsustainable rates and suffered serious population declines. Galapagos doves, hawks, and pintail ducks were killed in large numbers by residents of the islands for many years. While a few doves and ducks are still taken every year, populations of hawks in areas inhabited by humans are still prevented from recovery by human predation. Relatively recently vehicular traffic has become a serious source of mortality in several species of birds and at least one reptile. Automobiles on Santa Cruz Island account for several hundred dead birds a week and restoration efforts for land iguanas on Baltra Island are thwarted by mortality caused by buses.

Introductions

Some 600 species of vascular plant have so far been well-documented as introduced (excluding the 60 or so doubtful species that have been provisionally classed as native). The true figure is probably already over 1000. The introduction rate is often quoted to be exponentially increasing, with a recent rate of 10 per year (see Mauchamp 1997), but this is confounded by the artifact of detection rate, and may depend more on research effort than actual introduction rate. Regardless of the comparison of current to past rates of introduction, 600 species introduced


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since 1535 represents an overall rate of more than one species per year. The natural arrival-plus-establishment rate, as estimated by Porter (1983), is one species every 7000–12000 years. Therefore, since 1535 the introduction rate has been about 10,000 times the natural rate, and it has probably been even greater in recent years.

The native Galapagos flora is disharmonic in its species composition, as discussed in Chapter 3. This skewed representation of plant groups, relative to that on continental source areas, is caused by the filters to arrival and establishment represented by the sea barrier (only plants with appropriate dispersal mechanisms will arrive), survival barrier (only plants that can support the harsh Galapagos conditions), and establishment barrier (only plants that can successfully reproduce in Galapagos). Introduced species are selected differently. Man brings some species accidentally, side-stepping the need for efficient dispersal, and may transport their seeds directly to a favourable habitat. Humans bring far more species on purpose, for agriculture or ornament. In these cases, not only is the need for a dispersal mechanism circumvented, but also people assist the plant to survive, and provide enough seed for the immediate establishment of a viable population. If an introduction fails, people will often try again.

A further consequence of the species-poor and disharmonic characters of the flora is that the natural plant communities are highly susceptible to invasion, whether by better competitors (adapted to a species-rich continental environment), or by plants of life forms that are not represented in the native Galapagos community. An example is the susceptibility of the Fern-sedge Zone, which has no trees except the endemic treefern Cyathea weatherbyana, to invasion by introduced trees such as Quinine Cinchona pubescens and Guava Psidium guayava. Such species can completely change the structure of the plant community, shading out native species, as well as having other less obvious effects, such as root competition or reduction in suitable area for the growth of epiphytes.

For insects the probable native fauna comprises some 1500 species and a further 300 are classed as certainly or probably introduced (Peck et al. 1998). The natural arrival rate has been estimated at 1 species every 2000 years (Peck 1996), so the introduction rate since 1535 has been approximately 1 species every 1.6 years, or 1200 times the natural rate. In comparison to other oceanic island ecosystems, Galapagos has the lowest known proportion of introduced insect species. However, the introduction rate for invertebrates may actually be increasing, since insect introductions are accidental (there is no evidence of the intentional introduction of insects, even honey bees) and the volume of traffic to and between the islands is increasing. The introduction rate appears to have accelerated over the last three decades, correlating with the increase in human migration to the islands (Peck et al. 1998). In the first seven months of the implementation of an inspection and quarantine system in the archipelago, 33 insect species were detected on incoming organic produce although inspection activities were being carried out at a quarter of the capacity identified as necessary to implement the program in its entirety. At least three species had not been reported previously in Galapagos including the mangrove defoliator Thyrinteina arnobia (Causton et al. 2000). Peck et al. (1998) predict that the number of introduced species will double within the next ten years, especially since many species remain to be collected or identified. Numbers may already be much higher, as the urban and agricultural zones, probably the source of most introduced species, have not been thoroughly surveyed. This is also likely for other terrestrial arthropods: to date, 23 introduced species (principally diplopods, chilopods and isopods) have been introduced (Baert 2000).

The presence of introduced spiders, centipedes, millipedes, terrestrial isopods, earthworms and phytophagous insects is predictable in view of the soil and agricultural materials that have been moved to and around the islands. More surprising are seemingly fragile or physiologically maladapted introduced species such as terrestrial flatworms, and an Andean species of Onychophora in urban or agricultural areas.

Introduced invertebrates interfere in important ecological processes. Many such species are spreading and have the potential to invade a large range of habitats, displacing native species. At least seven introduced insects are serious threats to the flora and fauna of Galapagos. Two potential disease vectors have been introduced in recent years (Peck et al. 1998, Roque & Causton 1999): the mosquito Culex quinquefasciatus is elsewhere known to be a vector of bird malaria and other diseases including West Nile Virus; the Simulium blackfly is also a potential vector of bird diseases, while its larvae may be altering the nature of the stream ecosystems in the highlands of San Cristóbal. Two fire ant species (Wasmannia auropunctata and Solenopsis


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geminata) are aggressive predators and are seriously affecting native invertebrates by preying on them (Roque and Causton 1999; Roque et al 2000). Solenopsis is also known to interfere with the nesting behavior of birds and tortoises (Snell pers. obs.) Native invertebrates are also heavily preyed upon by two recently introduced predatory wasps (Polistes versicolor and Brachygastra lecheguana), which also remove prey insects from availability for native predatory species such as Darwin’s finches. Recent measures of prey volumes of Polistes show that it could eat more insect biomass in a year on some islands than all the Darwin’s finches combined on the same island (Parent 1999).

Native plant species are at risk from introduced phytophagous insects such as the cottony cushion scale insect Icerya purchasi, which attacks at least 61 native plant species, including Scalesia and Darwiniothamnus species, and is capable of causing mortality of the more susceptible plant species that are already debilitated (Causton 2001). Other newly introduced species, such as the whitefly Bemisia sp., are already having an impact on the ecosystem and could be a threat to endangered plant species such as the Scalesias.

There are also subtle cases of habitat change probably involving introduced invertebrate species. An altitudinal transect sampled for insects before and after the introduction of goats to Alcedo Volcano (Baert et al. 1999) showed changes in invertebrate diversity and composition. Phytophagous insects (associated with live plants) were replaced by xylophagous species (associated with dead wood), and high numbers of dry zone invertebrates, such as Scolopendron and scorpions, were found on the formerly more humid crater rim.

The situation with introduced vertebrates is quite different from that of plants and invertebrates. The number of introduced vertebrates (29+) is tiny compared to the number of introduced plants (600+) or invertebrates (300+). While the numbers of alien plants and invertebrates are large the relative numbers of them with great ecological impact appear less than that of vertebrates (see preceding paragraphs). Approximately 50 percent of the alien vertebrates (including humans) have apparently strong negative effects on indigenous biological diversity compare to the lesser percentages of obviously deleterious plants or invertebrates. Percentage of introduced plants that are naturalized is about 70% (depending on the definition of naturalized), and percentage of these that are causing problems or showing signs that they will is about 40% (or 30% of total introduced species).

Alien vertebrates contribute to population declines and extirpations of Galapagos organisms via direct effects such as herbivory, predation, parasitism, and potentially hybridization; as well as indirect effects such as competition, habitat alteration, and acting as vectors for alien pathogens. In many cases there are interactive effects and/or situations where a single alien species has different effects on different species. For instance – goats are herbivores of many indigenous plants, competitors of tortoises and land iguanas, and can promote erosion and reduction of ground cover that causes nest failure of tortoises.

Rates of colonization of alien vertebrates are less than those of plants and invertebrates. Assuming 24-29 species in 465 years, the rate is roughly one species every 16-20 years. The natural rate of colonization and speciation combined, the two processes leading to indigenous vertebrates, is approximately one species every 25,000 years. The natural rate of colonization only would be approximately one species every 50,000 years. Even though the rates for vertebrates are much lower than those of invertebrates the magnitude of the difference between alien and natural rates are surprisingly similar. Alien invertebrates are 1,600 times faster than natives and alien vertebrates are 1,250-1,550 times faster than natives. The relationship of alien plants to native plants is an order of magnitude greater than the relationships for animals (alien plants 10,000 times faster than natives).

Disease has been a major cause of extinctions in other archipelagos, for example of birds in Hawaii (Atkinson et al. 2000). In Galapagos a few species of domestic birds have been introduced - rock doves, chickens, turkeys – and in the captive populations of chickens there have been outbreaks of Marek’s disease and Newcastle disease. The incidence of diseases and parasites in Galapagos birds is largely unknown, but some initial research projects are under way. The continuing influx of potential disease vectors may be increasing rapidly the risk of alien diseases being able to establish populations in the islands. Some known vectors of disease have been introduced recently, thus increasing the risks of disease transmission. A considerable number of insect vectors of plant diseases have been introduced into Galapagos,


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such as white flies and aphids (Homoptera), yet we know nothing about plant pathogens in the archipelago. Still more disease vectors may have arrived but remain undocumented, for example, Diptera surveys have not been conducted since 1995. The Southern House Mosquito Culex quinquefasciatus was introduced in 1989. Adult females generally feed on birds and are suspected to be vectors of microfilariae in the blood of Galapagos penguins and flightless cormorants (B. Sinclair, unpublished). This species is also a known vector of avian malaria in other parts of the world and is thought to have contributed towards the devastating decline of endemic forest birds in Hawaii (Atkinson et al. 1993).

The blackfly, Simulium bipunctatum, is thought to have been introduced to San Cristobal Island in 1986 (Abedraabo et al, 1993) and has since been reported on other islands (Roque & Causton 1999). Whilst little is known about this species, others of the genus are vectors for the nematode worm, Onchocerca volvulus, which causes river blindness in humans (Shelley et al. 1997), and can also spread by mechanical transmission various animal pathogens, including avian trypanosomes, which infect both domestic and wild birds.

In short, introduced pathogens are an increasing threat, which urgently needs further research, as it does in other vulnerable locations (McCallum & Dobson 1995).

Once alien species arrive in the Galapagos they often continue to spread among the islands. Alien species of plants that have been in the archipelago for longer periods of time inhabit more islands than recent arrivals. However the relationship is complex because there are several species with long histories in the archipelago that inhabit only a few islands, but there are no species with a short history that inhabit many islands (Figure 5.2). The rate of spread among islands can be as great as a new island every two years. As with native species, large islands harbor significantly more alien species than small islands, but the relationship is weaker than with native species (Figure 5.2).

Alien Plants - Years inArchipelago & Distribution

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Figure 5.2. Distributional Patterns of Alien Species. The effect of years within the archipelago on dispersal is illustrated by regressing the number of islands inhabited by a species of alien p lant by the time elapsed since the first recorded observation of the species in Galapagos. The overall average is about an island every 90 years, the maximum rate is an island every two years. The species richness of aliens on islands is related to the size of islands, but not as strongly as with native species.


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Evolutionary Shifts

There are behavioral and morphological responses of some species that correlate with the presence of alien predators. Lava lizards on cat-free islets are significantly less wary and larger than on nearby large islets inhabited by cats (Stone et al. 1994, Snell et al. in prep). Cats have apparently functioned as agents of unnatural selection selecting for lizards that are warier. Cat predation does not actually select for small size, rather it selects for an early age at first reproduction, which in turn causes a small size.

It is likely that there have been other evolutionary shifts within Galapagos organisms caused by alien species that are as yet undetected.

Bottlenecks

As mentioned above, humans have brought about changes in the population structure of many species. The long-term evolutionary (and ecological) effects of such changes can at present only be guessed at, as in the example of change in size and age structure of timber trees.

Perhaps a more general effect is of genetic bottlenecks caused by severe reductions in population size, with or without changes in size or age structure. Many species have been reduced to a very few individuals, and the importance of the genetic variability that has been lost is completely unknown as is the extent of the loss. Species so affected include both plants (many species reduced to less than 1000 individuals, and several to less than 100) and animals, an example being the cacti and tortoises of Española Island.

Human population growth and distribution

In strictly biological and evolutionary terms, we humans are an alien species of vertebrate in the Galapagos, and our direct effects are previously described under extractive use of resources. However, many of our indirect effects appear correlated with our numbers and therefore patterns of our population growth are relevant in this section of threats to the Biological Diversity. In the earlier years of human colonization of the Galapagos, population growth was indistinguishable from that of continental Ecuador. Beginning in the late sixties and early seventies human population growth in the Galapagos accelerated (Figure 5.3). The population growth rate was well beyond that of the rest of the nation, and much of the increase appears related to immigration (see data in Fundación Natura et al. 2000). The immigration was probably caused by the real and perceived relative increase in the standard of living in Galapagos over that of much of Ecuador. The increase in the standard of living was largely due to economic opportunities provided by the increasingly successful tourist operations in the islands.

Figure 5.3. Human population growth in Galapagos.

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Artificially small populations and reduced habitats

As mentioned previously, populations of many organisms are severely reduced. Even if the causes of those reductions are removed, the fact that populations are smaller itself poses a distinct threat. Tiny populations are more susceptible to stochastic environmental variation. An example occurred with marine iguanas in 1980 on Gordon’s Rocks east of Santa Cruz. Gordon’s Rocks are usually inhabited by 100 or so marine iguanas and there is a small nesting area formed by a talus slope. In 1980 a locally heavy rainfall washed the talus slope into the sea, along with all of the eggs laid by marine iguanas that year. Thus that population of iguanas was probably unable to reproduce for many years until sufficient talus built up to replace the single nesting area. The population was not extirpated, but it certainly could have been if a few other factors had fallen into place.

Many plants have been reduced to such small populations that, not only is their gene-pool probably severely reduced, but other factors, usually unknown, may be preventing their regeneration following removal of the factor that caused their original decline. An example is the cactus Opuntia megasperma orientalis on Española Island. This species was extirpated from most of the island, and the small groups of plants left show little sign of regular recruitment. The factors affecting seed production, germination and survival, as well as the reasons for a lack of vegetative reproduction, are unknown. In this case, a reduced gene-pool and resultant lack of fertility does not seem to be the reason, since seed viability is high. Possible factors might include excessive predation on the few seeds produced, by the Large Cactus Finch Geospiza conirostris, but the reason could also be entirely natural, that regeneration normally only takes place in widely-spaced years when conditions are right. Long-term study may be required to settle this kind of question and manage the populations correctly to ensure their recovery.

There are many examples of artificially small populations in Galapagos reptiles. Populations of tortoises from Islas Pinta, Santiago, Santa Cruz, San Cristobal, and Isabela range between one to a few thousand individuals. Galapagos land iguanas from Baltra, Santa Cruz and southern Isabela ranged between 40 to 200 individuals prior to restoration efforts. The effects of these bottlenecks won’t be known for a long time, but most of these populations appear to have reasonable reproductive capacity so the damage may be low.

Pollution

Environmental pollution, that does not include live organisms, currently appears to be more of an aesthetic problem for Galapagos than a biodiversity conservation problem, but that situation could certainly change for the worse.

Patterns of climate change

Global climatic warming has received increasing attention of the past decade. On the short term it appears that the frequency and potentially the magnitude of El Niño phenomena may be increasing within the Galapagos (Snell and Rea 1999), reflecting a more general pattern within the Pacific Basin. Whether or not the ENSO oscillation itself is increasing, the prediction that it will be occurring on top of an elevated baseline sea temperature has great significance for Galapagos flora and fauna. Several populations of organisms have patterns of fluctuation that involve declines during El Niño events followed by recovery in the post-El Niño periods (marine iguanas, sea lions, fur seals, penguins and cormorants, for example). Furthermore it appears that the degree of population decline may be correlated with the magnitude of the El Niño event. Thus, if there is a potential for stronger El Niño events to happen more often, populations of sensitive species could suffer greater declines with shorter periods available for recovery. At some point recovery may not be possible and extirpations could result.

The potential effects of long-term global warming on the Galapagos could be significant, especially if warming occurred quickly. Rising sea levels would probably be of little importance to the natural systems, unless they occurred so quickly that beaches were lost. A warmer Galapagos would likely be a wetter Galapagos, which would probably favor many invasive species, whose current distributions appear partially limited by aridity. It is possible to speculate on many other potential problems, but the most general prediction is probably that, while


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climatic change may have little direct effect on much of the indigenous biodiversity of the Galapagos, its interactive effects with invasive species, patterns of habitat alteration, and exploitation of natural resources could be severe. Species adapted to cold conditions and having small populations, such as the penguin, could be particularly vulnerable to global warming.

Initiatives and opportunities for terrestrial conservation

The above summary of threats to terrestrial biodiversity is daunting but Galapagos presents important opportunities for addressing them. Foremost amongst the advantages enjoyed by Galapagos are the 40-year existence of the Galapagos National Park and the Charles Darwin Foundation, the Government of Ecuador’s commitment to conserving the Park (see Chapter 1), the capabilities of the Galapagos National Park Service, and the extraordinary international support that Galapagos can count on from UNESCO, WWF and others.

In response to the increasing gravity of the threats to biodiversity, the Government has taken some important actions in recent years. The most significant is the enactment of the Special Law for Galapagos (1998), which is discussed in Chapter 1 and Annex 1.2. Amongst its provisions are restrictions on migration to Galapagos, increased funding for the Galapagos National Park Service and local government authorities, and a framework for prevention and control of invasive species and environmental management in general. Migration control is already in effect, although with deficiencies in its application. So too is increased funding for GNPS: it receives 45% of Park entry fees to cover its marine and terrestrial responsibilities. However, for most conservation provisions of the Special Law to be effective, the Government needs to promulgate special regulations, which will specify how the various elements of the law are to be implemented. These regulations are in the process of being formulated.

In addition to legal measures, the Government has negotiated a US$ 18 million dollar grant from the Global Environment Facility via UNDP, to tackle the problem of invasive species in Galapagos. The project, which started in 2002, includes US$ 6 million for the eradication of goats from northern Isabela and US$ 5 million to capitalize an endowment fund. The Government has also obtained a loan from the Inter-American Development Bank, which includes US$ 1.8 million for infrastructure and equipment needed for the Galapagos quarantine inspection system.

On the ground, the Galapagos National Park Service and Charles Darwin Research Station have a strong track record of conservation achievements, most notably saving two populations of Galapagos tortoises and three populations of Galapagos land iguanas from extinction by the repatriation of captively reared juveniles. In all cases there is now reproduction by the repatriates occurring in the field. In the past five years GNPS and CDRS have scaled up their conservation, research and educational work. A joint team, known as Project Isabela, has eradicated goats from Pinta, has eradicated pigs from Santiago, is starting on the Santiago goats, and will, with specialist assistance in helicopter-based hunting, tackle the Isabela goats, when the GEF funds come through. Following this example, a joint program on introduced plants is also being developed and, taking advantage of the increased GNPS funding, will be eradicating several plant species that are still of restricted distribution, and controlling some of the worst weed species. In both plant and vertebrate work, the CDRS has substantially expanded its research programs, in support of the GNPS management efforts, although important topics have barely been touched on by either staff or visiting scientists, for example pathogens and parasites of vertebrates.

With regard to invertebrates, a field which has tended to be neglected in the past, the CDRS has over the past five years built up from scratch a permanent team and facilities. The team has been establishing a reference collection, collecting baseline data on the presence and distribution of native and introduced species, and studying some of the most noxious invasive species, notably the Australian cottony cushion scale, which affects over 60 species of native plants. The invertebrates team has also coordinated and supported the development of the Galapagos quarantine inspection system, known as SICGAL, which involves GNPS, the Ecuadorian Agricultural Health Service, CDRS and INGALA (the National Galapagos Institute) (Causton et al. 2000). The system started operations on a small scale in May 1999 and during 2000 intercepted 800 prohibited products on their way into Galapagos. As regulations are


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developed, personnel are trained and facilities are built, SICGAL will become one of the most important tools in the effort to exclude alien species from Galapagos.

Another important CDRS initiative of recent years, also in collaboration with the GNPS, has been to increase and re-orient investment in environmental education and cooperation with community groups. In addition to public dissemination of information and both formal and informal education, CDRS has been collaborating with key social groups, whose activities and attitudes affect the invasive species problem, for example farmers, traders and naturalist guides. Whilst there is no way to break completely the link between people and invasive species – every resident or visitor contributes to the risk of alien species introductions – the education and communication programs are vital to reduce the risk and to enhance the acceptability of new restrictions, such as prohibiting importation of certain goods, requiring that cargo be inspected, or controlling domestic pets.

On other kinds of threat, such as those posed by quarrying or exploitation of resources, less progress has been made and demand is increasing, as the growth and economic development of Galapagos proceeds apace, in contrast to the rest of Ecuador. Whilst the threats are potentially less serious than that of invasive species, they are significant sources of habitat destruction, especially for certain groups e.g. the snails, whose hillside habitats are being dug up for construction material. As in all fields, the GNPS capacity to manage these problems is increasing and the Special Law for Galapagos strengthens the legal framework for avoiding environmental damage.

SUMMARY: WHERE WE ARE NOW

The above discussions demonstrate that Galapagos in 1999 is far from the pristine state represented by 1535 but that, compared with other oceanic archipelagos, its biodiversity is still relatively intact. Key findings are:

• Most taxa are still extant but many are much reduced in numbers and declining.

• Most communities are still extant but many are much reduced in area; there are critical habitats that are not adequately represented in the park and continue to be heavily modified.

• Most endemic species and native communities are threatened in the short or long term, by factors that have been or are already in operation.

• Most plant communities and habitat types have suffered damage and this has been most severe in the humid highlands of the large islands.

• Primary threats are past habitat clearance, over-exploitation of a few species and, especially, introduced species.

• Alien species, many of which threaten the viability of native plants and animals, have been and continue to be introduced to the islands at rates that, for some taxa, are estimated to be up to 10,000 times the natural rate of establishment.

• Man has already caused dramatic changes that will continue to affect ecological and evolutionary processes into the future. International recognition and support, the 40-year history of the Galapagos National Park and CDF, and the strong partnership that exists between Park and Station, give Galapagos a great advantage in addressing the serious threats to biodiversity. Actions taken in the past 5 years have opened up important opportunities for addressing the threats to terrestrial biodiversity, in terms of the legal framework, financing, quarantine inspection systems and enhanced capabilities in conservation, research and education.


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REFERENCES

Abedraabo, S., Pont, F. le, Shelley, A.J. & Mouchet, J. 1993. Introduction et acclimatation d’une

simulie anthropophile dans l’île San Cristóbal, archipel des Galapagos, (Diptera, Simulidae). Bull. Soc. Ent. France, 98 (2).

Atkinson, C.T., Dusek, R.J. & Iko, W.M. 1993. Epidemic pox and malaria in native forest birds.

Hawaii's Forests and Wildlife 8(3): 10. Atkinson C.T., Dusek, R.J., Woods, W.L. & Iko, W.M. 2000. Pathogenicity of avian malaria in

experimentally-infected Hawaii Amakihi. Journal of Wildlife Diseases 36(2): 197-204.

Baert, L. 2000. Invertebrate research overview: 1. Terrestrial arthropods. Pp. 23-25. in Science for conservation in Galapagos. Ed. N. Sitwell. Bulletin de L’Institut Royal des Sciences Naturelles de Belgique, Vol. 70-Supplement.

Causton, C.E. 2001. Dossier on Rodolia cardinalis Mulsant (Coccinellidae: Cocinellinae), a potential biological control agent for the cottony cushion scale, Icerya purchasi Maskell (Margarodidae). Fundacion Charles Darwin.

Causton, C.E., Zapata, C.E. y Roque-Albelo, L. 2000. Alien arthropod species deterred from establishing in the Galápagos, but how many are entering undetected? Noticias de Galapagos. Vol 61.

Chambers, S.M. 1991. Biogeography of land snails. Pp. 307-326. In Galapagos Invertebrates: Taxonomy, biogeography and evolution in Darwin’s islands. Ed. M.J. James. Plenum Press, N.Y.

Cook, J., Howden, H. F. & Peck, S. B. 1995. The Galapagos Island’s genus Neorcytes Arrow (Coleoptera: Scarabaeidae: Dynastinae). Can. Entomol 127: 177-193.

Coppois, G. 1984. Distribution of Bulimulid land snails on the northern slope of Santa Cruz Island, Galapagos. Biol. J. Linn. Soc. 21: 217-227.

Coppois, G. 2000. Invertebrate research overview: 2. The endemic land snails. Pp. 27-29. in Science for conservation in Galapagos (ed. N. Sitwell). Bulletin de L’Institut Royal des Sciences Naturelles de Belgique, Vol. 70-Supplement.

Coppois, G. & Wells, S. 1987. Threatened Galapagos land snails. Oryx 21: 236-241.

Fundación Natura, The Nature Conservancy & WWF. 2000. Parque Nacional Galapagos - Dinamicas migratorias y sus efectos en el uso de los recursos naturales. Fundación Natura, Quito.

Groombridge, B. (ed.) 1992. Global biodiversity: status of the earth’s living resources. Chapman & Hall, London.

Groombridge, B. & Jenkins, M. (eds.) 1994. Biodiversity data sourcebook. World Conservation Press, Cambridge.

Lubin, Y. 1984. Changes in the native fauna of the Galapagos Islands following an invasion of the little red fire ant Wasmannia auropunctata. Biol. J. Linn. Soc. 21: 229-242.

Mauchamp, A. 1997. Threats from alien plant species in the Galápagos Islands. Conservation Biology 11, 260–263.

McCallum H., and A. Dobson. 1995. Detecting Disease and Parasite Threats to Endangered Species and Ecosystems. Trends In Ecology & Evolution 10(5): 190-194.

Parent, C. 2000. Life-cycle and ecological impact of Polistes versicolor versicolor (Olivier) (Hymenoptera: Vespidae), an introduced predatory wasp on the Galapagos Islands. Masters Thesis, Carleton University, Canada.


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Peck, S.B. 1996. Origin and development of an insect fauna on a remote archipelago: the Galápagos islands, Ecuador. Pp. 91–122 in The Origin and Evolution of Pacific Island Biotas (eds A. Keast & S. Miller). SPB Academic, Amsterdam.

Peck, S.B., Heraty, J., Landry, B. & Sinclair, B.J. 1998. Introduced insect fauna of an oceanic archipelago: the Galápagos Islands, Ecuador. American Entomologist 44: 218–237.

Porter, D.M. 1983. Vascular plants of the Galapagos: origins and dispersal. In Patterns of Evolution in Galapagos Organisms (eds R.I. Bowman, M. Berson & A.E. Leviton) pp. 33–96. American Association for the Advancement of Science, San Francisco.

Roque, L & Causton, C. E. 1999. “El Niño” and introduced insects in the Galápagos Islands: different dispersal strategies, similar effects. Noticias de Galápagos 60: 30-36.

Roque-Albelo, L., Causton, C.E. & Mieles, A. 2000. The ants of Marchena Island, twelve years after the introduction of the little fire ant, Wasmannia auropunctata. Noticias de Galapagos 61.

Shelley A.S., Lowry, S.A., Maia-Herzog, M. Luna Dias, A.P.A. & Moreas, M.A.P. 1997. Biosystemic studies on the Simuliidae (Diptera) of Amazonia onchoceriasis focus. Bull. Nat. Hist. Mus. Lond. (Ent.) 66(1): 1-121.

Snell, H.L., and S. Rea. 1999. El Niño 1997 – 1998 en Galápagos: ¿Se puede estimar 120 años de variaciones climáticos con estadisticas de 34? En: P. Ospina y E. Muñoz (eds) Informe Galápagos 1998 – 1999, pp 65-71. Fundación Natura, Quito, Ecuador.

Tye, A. 2002. Revision of the threat status of the endemic flora of Galapagos. Galapagos Report 6. Fundación Natura, Quito, Ecuador.

CDF/WWF Biodiversity Vision Chapter 6 – Terrestrial Biodiversity Vision

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CHAPTER 6 – PROJECTIONS FOR THE FUTURE: A TERRESTRIAL BIODIVERSITY VISION Principal Authors – H.L. Snell, A. Tye, C.E. Causton, G. Powell, E. Dinerstein, T. Allnutt and R. Bensted-Smith.

This chapter combines information from the previous chapters to construct a Biodiversity Vision for the Galapagos. We establish the recovery of all extant terrestrial biodiversity as the outer limit of possibility and ultimate long-term goal. We then describe two scenarios, one of which represents a set of ambitious goals that could be achieved in the coming 50-year period, if there are major improvements in conservation management of Galapagos; this is the Biodiversity Vision. The second scenario, which falls far short of this, is one set of likely outcomes based on the assumption that conservation and development continue along current lines for the next 50 years; this is the “business-as-usual” scenario. The projections represent our collective best judgment, based on knowledge of the ecosystem and ecological trends, combined with experience of what has happened in other island ecosystems, not on theoretical modelling. The two scenarios are described in terms of specific, measurable variables that could, if the necessary monitoring were to be carried out, be used to measure progress towards the 50-year vision and, beyond that, the ultimate long-term goal. However, in some cases there is little data and the estimates presented are quite speculative, giving only a general indication of expected trends.

6.1 Restoration Benchmark: The recovery of all extant terrestrial biodiversity of Galapagos

The Galapagos is probably the only remaining large oceanic archipelago in the world where: 1. We can still accurately quantify the distributions and conditions of terrestrial biodiversity prior

to human settlement, and 2. Virtually the full complement of biodiversity is still recoverable, and 3. It could be possible to restore the distributions of almost all terrestrial biodiversity to the

conditions prior to human settlement. Therefore, an approximation to the original pre-settlement state of Galapagos terrestrial biodiversity remains a valid, definable benchmark: an ultimate goal. To achieve it would take much more than 50 years and great advances in science and conservation, nevertheless we should not lose sight of it, as we struggle now to cope with the hordes of invading alien species. That was the feeling of the of participants in the Biodiversity Workshop, who described this ultimate goal as “the restoration of the populations and distributions of all extant native biodiversity and of natural ecological/evolutionary processes to the conditions prior to human settlement.” If this extremely ambitious goal were one day to be achieved, it would represent the pinnacle of accomplishment in conservation biology – the restoration of the biological nature of the Galapagos Islands almost to the conditions of 1534. PROJECTIONS FOR THE TERRESTRIAL BIOME Within the realm of what might be possible over the coming 50 years we have examined two scenarios: the first is that current trends continue (“Business as Usual”). Here we assume that current trends favorable to the preservation of Galapagos biological diversity continue as well as current trends that constitute threats to that diversity. This is not a dire “doomsday” scenario, because it recognizes the successes of recent years and improvements that we can reasonably expect in the future. Nevertheless, the fact remains that, even if no more alien species were to reach Galapagos, the processes of ecological change caused by the hundreds of plant and animal species that have already invaded the archipelago will be very difficult to stop and then reverse.


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The second scenario is what could happen if major improvements in the conservation of the Galapagos occur. It represents the vision for terrestrial biodiversity, based on an assumption of optimal conservation management over the coming 50 years. It aims for restoration of biodiversity but recognizes that, however good the conservation policies and practices, conservation takes time and faces many limitations. Thus, the vision is ambitious but attempts to stay within the realm of the potentially achievable. We present the two scenarios in a set of six tables. The first table (6.1) presents a core set of human activities that drive the threats to biodiversity described in previous chapters. Note that no specific assumption was made about global climate change. However, it is possible that an increase in frequency and intensity of El Niño phenomena could exacerbate introduced species problems. The second table (6.2) examines likely conditions under the two scenarios at the species level of criteria, the third table (6.3) at the communities/ecosystems level, the fourth table (6.4) at the landscape/habitat fragmentation level, the fifth table (6.5) at the ecological and evolutionary processes level, and the sixth table (6.6) examines specific conditions regarding alien species. Each table presents the 1999 condition and the two scenarios with brief explanations. The level of extinctions under “Business as Usual” presented in the Table seems low to some workshop participants and high to others. It is important to recall that the time frame is 50 years. In 50 years there will be tremendous population declines for many species and the situation will be dire. The projections estimate a substantial increase in the percentage of biological diversity recognized as endangered. However, many Galapagos organisms have adult life spans in excess of 50 or even 100 years. Thus the final act of extinction can take a tremendous amount of time (Galapagos tortoises on Pinzon Island appear to have failed to reproduce for over 100 years yet the population hasn’t changed noticeably in the last 30 years). The point is that while not many species will go extinct in the next 50 years, a substantial number will be set on the pathway toward eventual extinction in the future, if “Business as Usual” continues. As discussed in Chapter 5, disease is one factor that can cause rapid extinctions. The risk of extinctions due to introduced diseases is difficult to estimate and adds significantly to the uncertainty of extinction projections. The risk for Galapagos will be driven by both the rate of arrival of alien pathogens and the number and abundance of alien vector species, which may enable a newly arrived pathogen to establish. Another source of uncertainty in the projections is the changing global climate, particularly the likelihood of increasing intensity and frequency of the El Niño – La Niña cycle. In addition to its direct impacts, climate change may interact with other anthropogenic factors, such as invasion by alien species, in most cases exacerbating the problem but in a few cases amelioratiing it. The projections in this chapter assume that El Niño’s occur periodically, with a moderate increase in frequency and intensity due to global climate change.


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Table 6.1. Scenarios for Driving Forces Shaping the Future of Galapagos Terrestrial Biodiversity. Factor 1999 Status Business as Usual Vision Human population growth Total rate of increase 6%/yr

Result of immigration and natural increase of the resident population.

Total rate remains above 2% Current legislation could reduce immigration; Galapagos, like continental Ecuador, is undergoing a demographic shift towards reduced fecundity.

0% or negative Possible if immigration controls are enforced, the demographic shift occurs, economic tools such as incentives,and education and training improve substantially.

Distribution of human settlement

Settlement limited to Municipal, Agricultural, and Military Zones. Other infrastructure within GNP significant but still limited.

Park boundaries established by law, but airports, INGALA compounds, Tortuga Bay trail, Garrapatero road and Santa Cruz garbage dump are amongst the infrastructure inside GNP, and more proposals are in the pipeline.

Infrastructure and associated activities expand into areas previously protected within Galapagos National Park

Continuing pressure results in more development of roads and other infrastructure, perhaps using the controversial legal provision for 2% of inhabited islands to be allocated to INGALA. Around Bella Vista rural areas may be subdivided and urbanized. Southern Isabela settlements expand rapidly in density and impact. New solid waste disposal sites created in Park.

Municipal Areas Remain Stable and Agricultural Zones Shrink

Possible if none of the 2% is exploited and programs converting unused agricultural zone holdings to protected status are developed.

Transport, travel and tourism 5-7 cargo boats continuously delivering cargo from mainland to three islands. 2-3 plane loads of passengers and cargo per day. Increasing inter-island boat and air traffic. Increasing range and frequency of travel within inhabited islands. Numbers of vehicles and boats multiplying rapidly. Tourism approaching 70,000 visitors /year.

On all indicators numbers fluctuate, but general trend is an increase.

Volume of transport, travel and tourism continues to increase

Result of general trend occurring now. Isabela, hitherto more isolated and with fewer introduced species, may experience rapid increase. Numbers of vehicles and boats may be capped. Subsidies may be phased out. Cargo transport may become more orderly but volume increase. Development of locally based tourism may increase inter-island and within island travel.

Volume of transport, travel and tourism remains stable or declines slightly (especially cargo).

Possible with proper planning, regulation and use of incentives and disincentives in each of the three areas. Tourism planning will involve careful balance of factors: socio-economic, terrestrial conservation and marine conservation too.

Habitat clearance Overall amount low, but certain key zones nearly gone.

See Chapter 5. Some humid & highland habitats severely reduced.

Minimal Assumes no further clearing.

Negative Possible if no further clearing occurs and restorations of some humid & highland habitats occur, through purchase and incentives.


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Direct exploitation or other mortality on land caused by people e.g. road kills.

7 species across 10 to 15 populations exploited at non-sustainable levels1

See Chapter 5.

Unsustainable for 5 to 10 species1

Assumes that efforts to control and prevent unsustainable exploitation remain ineffective.

All exploitation of plants is sustainable; there is no exploitation of vertebrates.

Possible if current laws are enforced and penalties applied.

Frequency of introduction or dispersion of species relative to the volume of transport, travel and tourism.

Quarantine inspection data indicate high rate of undesirable transport of live organisms in cargo and by residents (accidental or deliberate).

Special Law for Galapagos includes measures to strengthen control of invasive species but little application so far, as regulations not yet produced. Quarantine inspection system initiated but only partially operational. Most people aware that introduced species are harmful but participation in combating the problem in urban and agricultural areas is low.

Significant reduction in rate of transport of undesirable organisms relative to volume of transport, travel and tourism.

Regulations promulgated. Donor investment in quarantine inspection for Galapagos is expected, but action to reform national body (SESA) is still needed. Tour operators’ voluntary environmental certification emphasises collaboration on introduced species problem. CDF and GNPS work with community generates increasing local participation in prevention, control and eradication.

Rate of transport of undesirable organisms relative to volume of transport, travel and tourism is reduced by about 2 orders of magnitude (depending on taxa).

Possible with full national commitment to the Galapagos inspection and monitoring system, combined with the evolution in Galapagos society of a culture of concern about the invasive species problem and active participation to combat it.

Availability of scientific information and research and management capabilities for prevention, control and eradication of introduced species.

Galapagos flora and fauna well studied but much more research needed to guide the control, eradication and restoration actions.

The GNPS/CDF partnership provides a strong management and research capability relative to other parks but insufficient for current scale of the invasives problem. GNPS financial resources increased by Special Law for Galapagos but still insufficient for major field programs.

Significant increase in GNPS/CDF management and research capabilities and hence in available scientific information, as well as in invasive species management action.

Substantial donor investment in invasive species problem is imminent. Endowment fund aims to help long-term sustainability.

Further increase in GNPS/CDF management and research capabilities and hence in available scientific information. Management methods developed and applied for many previously unmanageable alien species.

Possible if endowment fund continues to grow and government of Ecuador increases its commitment to finance Galapagos conservation.

1Matazarno (Piscidia carthagenensis), Tortoises from Southern Isabela (Geochelone guntheri, G. vicina), Galapagos Hawks, some populations of doves and ducks, land iguanas on Baltra.


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Table 6. 2. Projections of Galapagos Terrestrial Biodiversity – Species and Lower Levels. Criteria 1999 Status Business as Usual Vision Extirpations as % of total Diversity Species 1.5% of plants and vertebrates extinct.

Percentage of invertebrates unknown but probably higher.

Estimated using known number of vertebrates & plants as % of native species present in 1534. Probably an underestimate. Amongst invertebrates data are scant, but many Bulimulid snails have gone extinct, and perhaps some scarab beetles and terrestrial isopods (see Chapter 5).

Additional 3-15% of plants and vertebrates extinct.

Assumes some limited success in preventing extirpations, but assumes many species recognized as critically endangered will go extinct (Pinta Island Galapagos tortoises, Mangrove finch, Galapagos Petrel etc; at least 10 species of plants; invertebrate extinctions unpredictable, for lack of basic data, but many small island endemics and specialist feeders on endangered plants would be lost. The upper limit of the estimate is high, in recognition of the increasing but unpredictable threat presented by alien pathogens and diseases (these are also important in the following sections on extirpations).

No additional extirpations of plants and vertebrates. Some invertebrate extinctions inevitable.

Possible if new programs are initiated and new recovery methods discovered. Would require eradicating many alien species and preventing further spread of others, and improved restoration efforts. Also requires effective treatment of the increasing role of introduced pathogens and diseases.

Subspecies. Unconfirmed % extinct, probably 2 – 4%. Refers to taxonomically recognized subspecies. Percentage low because extinct forms unlikely to be named see “distinct populations”.

Additional 7 to 15%. Assumes some limited success in preventing subspecific extirpations, but assumes many subspecies currently endangered will go extinct (Vermilion Flycatchers of San Cristóbal for example)

Additional losses not exceeding 1%. Possible as explained above. However, even with new programs and recovery methods there will be some subspecies lost.

Distinct populations 2 % extinct. “Distinct populations” as a criterion removes the requirement of taxonomic recognition and considers the extirpation of complete populations. The value is estimated by combining values for vertebrates and plants and assuming for invertebrates, probably an underestimate. Amongst invertebrates some weevil populations are thought to have been extirpated.

Additional 7 to 15%. As above, assumes limited success, but many populations are assumed to go extinct (snakes from Floreana, land iguanas on Baltra, Opuntia on Floreana, snakes on Baltra, etc.)

Additional losses not exceeding 1%. Assumes that the increase can be held to 1% via active programs and new methods (see above).


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Criteria 1999 Status Business as Usual Vision Trends Towards Extinction % Diversity recognized as

Endangered or Critically Endangered (species & lower levels combined). See Annex 4.1 for definitions of “Endangered” and “Critically Endangered”. “Vulnerable” taxa are excluded.

24 % of endemic plants 50% of vertebrates Invertebrates unknown but 60% may be in decline.

Plants increase to 30% Vertebrates increase to 60% Invertebrates unknown but the 60% in decline may become endangered.

Assumes that the threats will continue to worsen sufficiently to endanger an additional 20% of vertebrates (the increase is only 10% because � 10 % of the currently recognized diversity will be extirpated).

Plants decrease to 10% overall Vertebrates decrease to 30% Invertebrates unknown but <30% in decline.

Possible as alien species are eradicated or their effects mitigated and other threats removed or lessened.

% Change in Abundance & Distribution from 1534 baseline.

Ranges from 0 to 90%. Greatest for humid zone plants and invertebrates, but also great for tortoises and several species of birds.

Will Range from 0 to 90%. Assumes that limited recovery will be offset by other declines.

Range from 0 to 20% in the National Park. 0 to 50% overall due to large numbers of species with nearly complete ranges outside of the Park.

Many at 0% or recovered to 0%. None reduced more than 20% within the Park..

% Species & populations with unstable population structure caused by anthropogenic factors.

70% overall. Refers to populations where reproductive success and or mortality rates indicate a declining population.

70% overall. As above, limited success will be offset by other declines.

<10% with unstable structure Possible with new programs, and greatly increased success with alien species.

Genetic & phenotypic variability compared to baseline.

Variation reduced in 60%. Represents an overall estimate. For phenotypic variation the coefficients of variation for representative traits are assumed reduced in 70 % or more of the diversity. Genetic variability may not be as reduced in many forms, but in some (Galapagos tortoises and petrels for example) it appears more reduced.

Variation reduced in 70%. Even if population levels begin to recover, the variability will take more time to recover. In some cases it is likely permanently lost.

Appropriate Coefficients of Variation reduced in <30%.

Assumes very quick recovery of population levels. In many organisms variability can be recovered if the populations did not remain low for long periods.


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Table 6. 3. Projections of Galapagos Terrestrial Biodiversity – Community and Ecosystem Levels, as represented by plant communities. Criteria 1999 Status Business as Usual Vision % Plant communities of which each of the distinct sub-communities (see Table 3.1) are represented in areas isolated from anthropogenic impacts.

8 communities have all sub-communities represented. 23 have some sub-communities represented. 5 have none of their sub-communities represented.

Most of the communities not represented in effectively protected areas are of the humid zone, see Table 3.1 and chapter 5 for details.

Same as for current status. There will be little improvement because the habitats without representation are those within or near the agriculture zones.

32 communities have all sub-communities represented. 4 have some sub-communities represented. 0 have none of their sub-communities represented.

Possible only if specific areas within the agriculture zones were to become protected areas and if great advances made in controlling invasive species. The 4 communities in the “some” category are, in the arid zone, Opuntia-Scalesia open scrub, and three in the humid zone: Scalesia zone, Brown zone and tree-fern groves.

% humid highland habitat uncleared compared to baseline.

Ranges from 7-100% :- Santa Cruz – 26% Floreana – 85% Cristóbal – 7% Isabela (all) – 92% Isabela (Sierra Negra) – 86% Santiago – 99% Pinta – 100% Fernandina – 100%

The large reductions are caused by the overlap of the agriculture zone with highland habitats. Note that arid and littoral zone habitats, not covered by this indicator, are almost entirely uncleared.

Ranges from 5 -100% :- Santa Cruz – 24% Floreana – 80% Cristóbal – 5% Isabela (all) – 90% Isabela (Sierra Negra) – 80% Santiago – 99% Pinta – 100% Fernandina – 100%

Little improvement due to extent of agricultural zones. Some further clearance for development purposes.

Ranges from 30 -100% :- Santa Cruz – 50% Floreana – 85% Cristóbal – 30% Isabela (all) – 95% Isabela (Sierra Negra) – 90% Santiago – 99% Pinta – 100% Fernandina – 100%

Possible with restoration of agricultural land to natural habitat. However, complete recovery will remain impossible due to required agriculture.

% humid highlands that is uncleared and has species composition not significantly altered relative to baseline

Ranges from <1-100% :- Santa Cruz – <5% Floreana – 10-20% Cristóbal – <1% Isabela (all) – 30-40%

Ranges from <1-70% :- Santa Cruz – <1% Floreana – <5% Cristóbal – <1% Isabela (all) – 10-20%

Ranges from 10 -100% :- Santa Cruz – 10-20% Floreana – 20-30% Cristóbal – 10-20% Isabela (all) – 50-60%


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Criteria 1999 Status Business as Usual Vision Isabela (Sierra Negra) – 20-30% Santiago – 5-10% Pinta – 5-10% Fernandina – 100%

This is more restrictive than the above indicator, as it takes account of the alteration of natural communities by alien species. These are very rough estimates, as no reliable data are available.

Isabela (Sierra Negra) – 5-10% Santiago – 5-10% Pinta – 5-10% Fernandina – 60-80%

Assumes that eradication of alien species will be limited. Fernandina will be affected by invasive alien species crossing over from Isabela, as well as direct anthropogenic introductions. In Pinta the goats have gone but the restoration of natural communities is impossible without the restoration of the tortoise population, currently down to just one individual (Lonesome George).

Isabela (Sierra Negra) – 40-50% Santiago – 70-80% Pinta – 90-100% Fernandina – 90-100%

Possible if alien species removed from great areas, with intensive monitoring and rapid response. Santiago assumes pig and goat are eradicated. Pinta assumes an appropriate tortoise population is re-established. However universal eradication is impossible, thus some communities will remain with altered species composition, especially on the inhabited islands, where many invasive species, that are very difficult to control, are established and spreading, and further arrivals are inevitable.


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Table 6. 4. Projections of Galapagos Terrestrial Biodiversity – Landscape Level.

Criteria 1999 Status Business as Usual Vision Increase in fragmentation of key habitat types from baseline

0 – 20x A few highland habitats are extremely fragmented. These values are subjective estimates that require confirmation, but several habitats seem at least twenty times as fragmented as we assume they were at the baseline. Some mangrove habitats are also fragmented around settlements.

0 – 20x Without large scale conversion of the highlands there can be little reduction in fragmentation.

0 – 5x Possible if areas between remaining fragments can be strategically restored. Small isolated blocks could be combined into larger tracts. We assume that this would still result in a remaining fivefold increase in the worst cases of fragmentation over the baseline.

Increase in isolation of fragments

0 – 50x As habitat fragments are lost, the distance between remaining fragments increases, decreasing the potential of connections.

0 – 50x As above.

0 – 10x Possible if missing fragments between existing ones can be restored, or if the size of fragments canbe increased by restoration.

% of archipelago’s area composed of “near pristine” islands, defined as islands with no alien vertebrates, < 5 alien plants, no unnatural extirpations and no large-scale anthropogenic changes (see chapter 4).

10% Only 0.007% of the area of Galapagos is composed of completely pristine islands (the largest is Caldwell Island, 22 ha). The largest “near pristine” island is Fernandina (64,000 ha), assuming that the anis seen there in 1998 have failed to establish (they have not recently been observed). The next largest is Marchena Island, 13,000 ha.

3% While the number of near pristine islands will increase as restoration successes occur, all apart from Fernandina will be small islands and therefore not really affect the total area very much. Fernandina itself is unlikely to remain near pristine for 50 years under business-as-usual.

11% This is possible with the restoration of some relatively very large islands and the maintenance of Fernandina as near pristine. However, due to the tremendous size of islands like Isabela, Santa Cruz, San Cristóbal, and Santiago, all with more than 100 alien species, this percentage is capped at around 11%.


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Table 6. 5. Projections of Galapagos Terrestrial Biodiversity – Ecological & Evolutionary Processes. Criteria 1999 Status Business as Usual Vision

Ecological Processes Biotic Interactions (% of communities with altered patterns of herbivory, predation, or competition).

90 % Alien species are nearly ubiquitous in Galapagos, therefore the vast majority of communities experience some alteration in at least one form of biotic interaction.

80% With moderate success at eradicating alien species some communities will be returned to completely natural sets of biotic interactions.

50% Possible if large-scale eradication of alien species is realized.

% communities with frequent unnatural disturbances (other than those caused by alien species).

10% This is an area that must be quantified in the future. We have little data on unnatural disturbances but there have been some large fuel spills and anthropogenic fires.

10% We assume that current trends will not control these disturbances enough to reduce their frequency or distribution.

0 % Possible with effective control of human activity to prevent fires and spills.

Evolutionary Processes % of Species experiencing unnatural selective pressures.

50% Measures micoevolutionary processes. The altered biotic interactions mentioned above constitute unnatural selective pressures for some species. Examples are the effects of cat depredations on size & behavior in lava lizards and snakes.

50% Assumes that the status quo will be maintained. Moderate successes are predicted to be offset by losses.

40% Possible with large-scale eradication & mitigation of alien species.

% Species experiencing altered rates of gene flow (result of reduced or increased isolation).

10% This is caused by an alteration of isolation. The movement of native species among islands breaks down isolation and may prevent the process of speciation. Habitat fragmentation increases isolation and may prevent local adaptation by reducing genetic variability. Again, this is an area with very little data at present.

10% Effective quarantine may decrease the increased gene flow, but increased amounts of material movement are likely to offset the gains. Quarantine may be less effective at preventing movement of native species due to potential biases in inspectors.

2% Potentially possible with strict control of movement and quarantine, combined with reduced fragmentation of habitat.


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Table 6. 6. Projections of Galapagos Terrestrial Biodiversity – Alien Species. Criteria 1999 Status Business as Usual Vision Number of alien species that arrive at and colonize Galapagos per year

10 plants, 10 Invertebrates, 0.5 vertebrates

These are average rates of arrival and colonization over recent years.

10 plants, 10 invertebrates, 0.5 vertebrates

Assumes that increased efficiency of quarantine will be offset by increased amount of material moved.

Near-zero for all With effective quarantine and reduction in amount of material moved, it should be possible to reduce introduction rates by two orders of magnitude or more, depending on the taxa, albeit never entirely preventing new introductions.

Increase in distribution among islands of alien species already established in Galapagos in 1999. This excludes the spread or eradication of species that become established in Galapagos after 1999.

Index of current distribution1: Plants 1,200, Vertebrates 81.

i.e. the sum of the numbers of alien species on each of the 17 larger islands in 1999 is approximately 1,200 for plants and 81 for vertebrates.

Pre-1999 inter-island colonization has varied greatly, with certain species colonizing up to 27 of the 128 islands, in addition to the single island they are presumed to have initially colonized.

There have also been pre-1999 eradication successes on specific islands, particularly for vertebrates.

Colonization index2: Plants 240, Vertebrates 65.

i.e. the sum of the numbers of new colonizations of each island (within the list of 17 larger islands) by alien species, that were already established in Galapagos in 1999, is projected to be 240 for plants and 65 for vertebrates. Eradication index2 Plants 20, Vertebrates 14.

i.e. the sum of the number of species, already present in 1999, that are subsequently eradicated from each of the 17 larger islands is projected to be 20 for plants and 14 for vertebrates. These projections assume that limited eradications will be offset by increased spread of alien species due to increased inter-island movement. It is projected that some species may spread to as many as 30 islands, in addition to the island where they first established.

Colonization Index2: Plants 90, Vertebrates 14.

Eradication Index2: Plants 40, Vertebrates 50.

Possible to achieve these improvements with aggressive programs of eradication and control of movement, although quarantine can never be 100% effective in preventing new introductions and spread. Most plant species, once they are well established, are very difficult to eradicate. No distribution index, like that in the 1999 column, is estimated for the Vision or Business-as-usual. However, CDF plans in 2002/3 to collect more complete data, in order to verify the baseline situation, and thereafter to use this indicator (and the detailed island-by-island data used to calculate it) to track and project the overall number and distribution of alien species in Galapagos.


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Criteria 1999 Status Business as Usual Vision % of alien species eradicated from Galapagos (of those that have ever become established in Galapagos).

0.4% plants (two species), 0 invertebrates, 0 vertebrates

This criterion is eradication of the alien species from Galapagos, not just from a particular island or region. We’ve not yet been successful at this.

5% plants, 1% invertebrates, 16% vertebrates

Assumes complete eradication of at least 30 species of plants, a few invertebrates, and 4 vertebrates (anis, burros, rock doves, possibly green iguanas).

10% plants, 5% invertebrates, 80% vertebrates.

Possible with aggressive programs of eradication and simultaneous programs to promote public support.

% Species currently endangered by aliens that will be protected by mitigation

10% In some cases, it is not currently possible to eradicate alien species with serious impacts on native species. Those impacts can be mitigated by controlling the numbers of the alien species or by protecting the native species. A few species are currently protected (Pinzon tortoises, Galapagos Petrels, Linum cratericola, Scalesia atractyloides etc.).

30% Research in effective mitigation seems moderately promising for several species of native organisms (snakes, Scalesia, etc).

100% Certainly possible if commitment is made for funding and implementation. Successful mitigation is only limited by effort.

1 The current distribution index is the sum of occurrences of alien vertebrates and alien plants on the 17 islands larger than 150 ha in 1999. This is provided as a basis of comparison for the remaining two columns. In the case of large islands that have not recently been surveyed by botanists, we have estimated the 1999 numbers of plant species on the basis of prior data plus general observations. A full set of data for these 17 islands should be available by about 2004 and, together with the vertebrate data, will provide a valuable baseline. The possibility of weighting this indicator by island area has been considered, as has the possibility of weighting it by the number of native species that each island possesses. All three options have their merits in summarising in a single figure the extent of the alien species problem in Galapagos. The 17 islands included are Isla Isabela, Isla Santa Cruz, Isla Fernandina, Isla Santiago, Isla San Cristóbal, Isla Floreana, Isla Marchena, Isla Española, Isla Pinta, Isla Baltra, Isla Santa Fé, Isla Pinzón, Isla Genovesa, Isla Rabida, Isla Seymour Norte, Isla Darwin, and Isla Wolf.

2 The Colonization Index is the estimated future sum of colonizations of 17 islands1 by alien species present in the archipelago but absent from the particular island in 1999. The Eradication Index is the estimated future sum of eradications from one of the 17 islands of alien species that were present on one of the 17 islands in 1999. Both indices are presented separately for plants and vertebrates.

CDF/WWF Biodiversity Vision Chapter 7 – Outstanding Marine Features

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CHAPTER 7 - OUTSTANDING MARINE FEATURES OF GALAPAGOS Principal Authors - R.H. Bustamante, G.M. Wellington, G.M. Branch, G.J. Edgar, P. Martinez, F. Rivera, F. Smith and J. Witman Location and environmental setting: The Galapagos Archipelago is located in the equatorial eastern sector of the Pacific Ocean, ca.1000 km off the coast of Ecuador, South America, between 01°40´N-01°25´S and 89°15´W-92°00´W. The archipelago consists of some 130 large and small islands and islets, with the largest island – Isabela – being 130 km x 40 km. Combining all the islands and using a map scale of 1:600,000, the total surface area is about 50,130 km2 and the coastline ca. 1,800 km (Snell et al.1995; Bustamante & Vinueza unpublished data). The islands represent the tops of relatively young emergent volcanoes that rose from the sea between 5 and 9 million years ago (Christie et al. 1992), and they comprise a relatively shallow (<200m) Galapagos Platform surrounded by deep waters (+1,000-4,000m) (Fig. 7.1). The location of the Galapagos Archipelago, at the confluence of warm and cold surface currents and deep, cold, upwelling waters, has led to complex marine and coastal ecosystems that are presently poorly understood (Colinvaux 1972, Wellington 1984, James 1991).

Figure 7.1 Bathymetric contour of the Galapagos platform. The color bar indicates depth in meters (Data and figure developed by Dr. Bill Chadwick, Hatfield Marine Science Center Oregon State University/NOAA, http://newport.pmel.noaa.gov/~chadwick/) The geographical setting provides the islands with a unique oceanographic environment, comprising a tropical archipelago situated between major ocean currents and under persistent upwelling conditions (Houvenaghel 1984, Chavez and Brusca 1991). Figure 7.2 depicts the general patterns of major ocean currents. From the north, warm (26-29°C), nutrient-poor waters flow from northeast to southwest, with a strong seasonal influx during the Austral summer (January to March). Over the years this current has brought most of the tropical elements that contribute to Galapagos marine biodiversity today. Flowing from the southwest, a major influx of cold (13-18°C), nutrient-rich waters reach the islands from the Chile-Peru current system (an extension of the Humboldt Current, in blue in Figure 7.2). This flow brings the majority of cold-adapted species that inhabit the Galapagos. The confluence of these two major north and south influxes, in addition to prevailing winds and the Coriolis effect, drives most of the surface waters towards the west. This gives rise to the South Equatorial Current (shown in white in Figure 7.2), which has been proposed as the major “vehicle” bringing species from mainland


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Ecuador to the islands. Flowing in the opposite direction from west to east, the Equatorial Countercurrent or Cromwell Current (in blue, Fig. 7.2) – a deep, subsurface cold and nutrient-rich current – hits the Galapagos Platform and disperses through the archipelago, creating persistent nutrient-rich upwelling cells off the western shores of most islands (in green, Fig. 7.2).

Figure 7.2 Schematic depiction of the major currents and water movement around the Galapagos Islands. Blue arrows indicate the main cold-water flows and red arrows the warm waters. In green are depicted the localized upwelling cells of high productivity (Art produced and designed by Mats Wedin, CDRS Communication & Education Department). Galapagos biogeography: Cold and warm influxes interacting in a highly dynamic environment, and acting over an evolutionary time period, have produced several discrete biogeographic zones that are separated by very short geographical distances (Abbot 1966, Harris 1969, Glynn and Wellington 1983, Bustamante et al. 1995). Between three and five major biogeographic units have been proposed for the archipelago; however, the number and boundaries still require further investigation (Harris 1969, Jennings and Brierly 1994, Wellington et al. 2000). Figure 7.3 depicts the five biogeographic units proposed by Harris (1969). Despite being proposed more than 30 years ago, this model of biogeographic division still appears valid. Data on benthic invertebrates and seaweeds have shown that at least three of the five proposed zones depicted in Fig. 7.3 - the cold, warm and mixed temperate biogeographic units - can be identified with distinctive and unique biota that have colonized the islands from four different sources (McCosker and Rosenblatt 1984, Kay 1991). These are the Central Pacific, the Ecuadorian mainland, the Chile-Peru region, and the Panama-Caribbean region (James. 1991). Recent quantitative studies by CDRS in over 250 coastal sites around the archipelago have highlighted the distinctive character of the western part of the archipelago and may lead to some refinement of Harris’s five units (G.J.Edgar pers. comm.).


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Figure 7.3 Diagram representing the main five biogeographic units proposed by Harris (1969). The outer lines represent the boundaries of the Galapagos Marine Reserve (GMR). The heterogeneous environment of the Galapagos Islands is also compounded by large-scale environmental changes, notably the El Niño-Southern Oscillation (ENSO). During El Niño years a reversal occurs in the normal atmospheric pressure gradient across the mid-Pacific Ocean, with the normal pattern of high pressure in the western Pacific disrupted (Allan et al. 1996). Atmospheric pressure during an El Niño is anomalously high over the Galapagos Islands, depressing the sea level and causing warm currents to flow into the region. Much of the spatial variability in ocean climate through the region disappears and water temperatures maintain a consistent level at around 29 oC. Associated with these high temperatures are clear water, a pronounced depression in the depth of the thermocline (down to 300 m), low nutrient concentrations, low phytoplankton productivity, and low ecosystem productivity. El Niño events are massive in both time (months to years) and space (hundreds of kilometers), and have huge impacts on marine ecosystems in the Galapagos region (Glynn 1994). They are also frequently followed, often in the subsequent year, by prolonged periods of anomalously low atmospheric pressure, high sea level and the intrusion of cold-water – a situation known as La Niña. Thus, Galapagos marine environments are extremely variable not only in space but also over time – few places experience such dramatic changes in sea surface temperature in areas less than 10 km apart (from west to north), from year to year, or even from one day to the next (Bustamante et al. unpublished data). The Galapagos marine environment thus imposes a strong, selective pressure for the ability of the biota either to survive extreme environmental fluctuations or to rebound quickly after a severe population reduction. This highly dynamic system with numerous species at their tolerance limits makes trends in Galapagos marine biodiversity difficult to monitor and predict, and may also make the ecosystem particularly vulnerable to additional human-induced stresses like intense fishing, habitat modification, or severe and persistent pollution.


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Habitat diversity: Two major complexes of marine habitats can be found: the deep open ocean and shallow coastal shores. Table 7.1 lists the major habitats identified in the Galapagos. Most of the coastal shores of the Galapagos consist of hard-bottom, consolidated sloping lava fields. More than 90% of all shallow benthic habitats correspond to lava reefs, with interspersed sandy pocket beaches composed of both biogenic material (white and brown sand mainly from corals and echinoid exoskeletons) and pulverized lava (black sand). These habitats result from volcanic and bioerosion activities that Galapagos has experienced over the years. Lava reefs are present at all islands and are interspersed with other habitats such as vertical walls, sandy beaches, and mangroves. Of all habitats, coral reefs are the rarest, as they are restricted to a few patches of several hundred square meters. Coral reefs are rapidly declining due to ENSO stresses and coral grazing by dense populations of sea urchins and fish (Glynn et al. 1979, Glynn 1990, 1994). Mangroves are also highly localized in the Galapagos (Farnsworth & Ellison 1997), and are restricted to a few major sites. Four species of mangrove trees are present; Avicennia germinans, Conocarpus erectus, Laguncularia racemes and Rhizophora mangle. Little is known about the mangrove communities and biodiversity patterns. The tidal range of the Galapagos is semidiurnal, with two low tides and two high tides each day (INOCAR, 2000). The tidal range is about 2.5 m, creating a total area of around 41 km2 of intertidal habitat. Similar to most tropical intertidal habitats, the black lava rocks reach high temperatures during daytime low tides, restricting most of the mid-to-high shore biodiversity to cryptic habitats (e.g. under boulders and in crevices). In contrast, in the low-shore the intertidal biodiversity is abundant and conspicuous. To date, intertidal communities of the Galapagos Islands have not been adequately assessed. Substantial areas of coastal lagoons and wetland habitats only occur in southern Isabela. However, small tidal lagoons are present in most of the large- and medium-sized islands. The numerous, complex lagoons of southern Isabela were formed by rainwater running off the slopes of the southern volcanoes and collecting in the lowlands. In this area are found the nesting grounds of flamingoes (Phoenicopterus ruber), the feeding and nesting grounds of resident and migrant waders (e.g. Galapagos crake, plovers, wandering tattler, American oystercatcher, whimbrel, common gallinule, and ruddy turnstones), the nesting beaches of green sea turtles (Chelonia midas), and endemic species that occur in brackish habitats. These habitats remain largely unexplored, as is shown, for example, by the recent discovery of the brackish water eel Anguilla marmolata, with a length of 1.6 m (McCosker et al. in prep.). With the development of the Geographical Information System (GIS) for Galapagos, habitat distributions are now being assessed. In the deep waters around Galapagos, six major habitat types can be found – upwelling zones, seamounts, pelagic waters, Galapagos shelf and slope, abyssal plains and hydrothermal vents. Although these habitats overlap to a certain extent, the species associated with these habitat types are generally distinctive (Cairns 1986, Monniot & Monniot 1989). The first habitat type comprises areas where upwelling occurs. Elsewhere in the world, where this habitat has been better studied, cold upwelling systems have been found to harbor unique sets of plants and animals, hence this pattern can be expected to hold for Galapagos, particularly the west coast (Longhurst 1985, Cushing 1989, Emanuel et al. 1992, Bustamante et al. 1995). The second habitat type, represented by the shallow seamounts (between 100-300m depth), is thought to be formed by submerged volcanoes that create topographically localized oceanographic conditions. These areas are normally associated with rich faunal communities (Genin et al. 1986). Much of the land-based fauna (e.g. seabirds, sea lions) feeds around these seamounts, which are mostly located west and south of the Galapagos platform. Shallow seamounts also attract considerable fishing activity. The truly open pelagic system is the most abundant and extensive habitat off the Galapagos Islands. It harbors several species of cetaceans and other higher vertebrates, numerous species of open-water fish, and planktonic communities. The Galapagos Shelf and offshore slopes consist of the benthic rocky habitat of the Galapagos platform, which connects most of the islands at depths between 100-300 m. The abyssal plains, at depths greater than 1,000 m, are the largest and least explored of the ocean’s habitats world-wide. Virtually no research or biodiversity inventories have been conducted for abyssal habitats around the Galapagos Islands. Notably, the hydrothermal vents of the Galapagos spreading center were the first ones to be described and presented to science (Lonsdale 1977). Similar habitats are expected to occur near the seeps in the mantle crust around the location of the western Galapagos hot spot (D. Geist, pers. comm.).


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Table 7.1 Main coastal and open water habitat found in the Galapagos Islands

Coastal & Shallow (<100m) Offshore & Deep waters (>100m)

Rocky Intertidal Upwelling Zones

Rocky Subtidal Seamounts (“Bajos”)

Sandy Beaches Pelagic Ocean

Vertical Walls Galapagos Shelf and Slopes (>100-300m)

Coral Reefs Abyssal Plains

Mangroves Hydrothermal Vents

Coastal Lagoons

Species Diversity: In addition to its particular historic and geographical position, the isolation of the Galapagos Islands has led to a high proportion of endemic marine species. To date, 2909 marine species have been recorded (Table 7.2). Because the marine ecosystems of Galapagos are less isolated in ecological terms than those on land, they tend to have low levels of endemism compared to terrestrial ecosystems. The proportion of marine endemism for Galapagos, averaging about 21% per taxonomic group and 18.2% as a total (Table 7.2), is high compared to other marine islands and archipelagoes, which range between 0-20%. It is also important to note that several prominent Galapagos endemic vertebrates, such as the marine iguana (the world's only sea-going lizard), the flightless cormorant and the Galapagos penguin, depend on the sea. Table 7.2 depicts the numbers of species per major taxonomic and functional group, compiled from available literature. The islands are noted for their spectacular creatures: sharks, whale sharks, cetaceans and manta rays, as well as consistently-abundant, commercially-valuable pelagic fish such as bill fish and tuna. The number of known species of marine plants and animals for the Galapagos has been constantly increasing since 1990, due to new taxonomic and biodiversity inventories. Recent explorations of deep-sea communities, especially invertebrates and fish, are yielding new additions to science and to Galapagos marine biodiversity. In terms of species richness, the Galapagos marine ecosystem is in the intermediate-to-high range, in comparison with other insular systems (Hawaii, Marquesas, etc.), despite its biological communities being less well studied than many archipelagos. Table 7.2. Number of marine species per trophic or functional group recorded in the Galapagos Islands, indicating level of endemism, relative abundance in relation to other oceanic islands, and level of study.

Groups # Species

# Endemics

% Richness** Level of Study

Mammals 24 2 8.3 High Good Algae ↑ 333 130* 39.0 High Poor Marine Birds 19 5* 26.3 High Good Fish ↑ 447 51* 11.4 Intermediate Moderate Soft bottom communities

390 ? ? High Poor

Polychaeta 192 50 26.0 Intermediate Poor Amphipods 50 19 38.0 Intermediate Good Brachyurans ↑ 120 23 19.2 Intermediate Poor Caridea & Stenopods ↑ 65 10 15.4 High Poor Porcelain crabs ↑ 12 1 8.3 Low Good Barnacles ↑ 18 4 22.2 Low Good (cont)


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Groups # Species

# Endemics

% Richness**

Level of Study

Mollusks ↑ 800 141 17.6 Low Poor Opisthobranchs ↑ 49 18 36.7 Low Poor Echinoderms ↑ 200 34 17.0 High Moderate Bryozoans 184 34 18.5 High Poor Gorgonians 12 8 66.7 Low Poor Corals 44 20 45.5 Low Moderate

Totals 2909 ♣♣ 531 18.2% ↑ Groups that have new records or species not included in the figures ♣♣ Not including recent deep-water surveys * Including island endemic species ** Relative to other Pacific insular areas The total reported species richness of individual islands in Galapagos is strongly related to island size (Fig. 7.4). While the relationship for literature and taxonomic records partly reflects sampling effort at the different islands – i.e. too few species for large under-sampled islands – quantitatively surveyed islands showed that a similar but less dramatic trend also exists (Fig. 7.4). One hypothesis explaining these results is that large islands are likely to have a greater diversity and abundance of habitats, and consequently more species are able to occupy distinct ecological niches. Different parts of larger islands could also be affected by the various local ocean currents, allowing more than one biogeographic unit to exist per island. For example, large islands like Isabela or islands that are situated at the boundary of several biogeographic units (see Fig. 7.3) will exhibit high species richness compared with islands situated in one biogeographic unit, due to more varied sources of colonization over time. This apparent pattern is consistent with the island biogeography theory of MacArthur and Wilson (1967); nevertheless, the relationship between species distribution and diversity needs further study for Galapagos marine ecosystems.

Spatial Pattern of Galapagos Marine Biodiversity

1

10

100

1000

10000

1 10 100 1000 10000 100000 1000000

Island Area (ha)

#Spe

cies

Literature+C. Hickman Records # Sps Surveys

Known species richness

Figure 7.4. Relationship between island size (in hectares) and number of species present or recorded on each island. Dots represent the number of recorded marine species (invertebrates, fish, and algae) per island from both literature and taxonomic records (solid circles) and recent quantitative surveys (hollow circles). Quantitative taxonomic surveys have yielded interesting marine biodiversity distribution patterns. Of the five biogeographic regions proposed by Harris (1969), recent data show that the most species-rich regions are the central to southern of the Archipelago (Fig. 7.5). This pattern is consistent not only with recent surveys but also with pre-existing records of the marine biodiversity listed in Table 7.2. The northeastern and central lower-north regions exhibited the lowest diversity of faunal and floral species. This pattern may be deceptive, however, since these coarse approximations of species richness do not necessarily reflect patterns within


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taxonomic groups, which may have colonized from different sources. In summary, the central, southern and western areas are the focal point of the new arrivals of all species from three different sources: the central East Pacific; the South American mainland; and the Chile-Peru Humboldt system (see Fig. 7.2). In contrast, the northeastern regions are primarily influenced by the tropical component (Fig. 7.5), in which some particular groups are more species rich (e.g. corals and fish). The northern islands harbor the highest reef fish species richness, while the west harbors the lowest. Conversely, for marine algae, the western islands (Fernandina and western Isabela) have the most diverse and abundant marine flora within the Galapagos Islands (Silva 1964, Wellington 1974), reflecting the higher productivity there due to upwelling. Although comprehensive assessments of diversity are being conducted in subtidal regions of the GMR, the pace of diversity change is virtually unknown and needs to be investigated by long-term monitoring. A recent study showed that the rate of diversity increase in sessile invertebrate communities at some sites in the GMR can be unusually rapid (Witman and Smith, in press). For example, the species richness of epifaunal invertebrates in rock wall habitats doubled between 1999 and 2000 at a site in the central sector of the Archipelago (Witman and Smith, in press).

Figure 7.5. Total number of shallow benthic species per biogeographic region in both literature records and taxonomic surveys. Genetic diversity and evolutionary processes: Endemism In contrast to terrestrial environments, where distances between islands can pose a significant barrier to gene flow, most marine organisms in Galapagos waters produce long-lived pelagic larvae or spores capable of widespread dispersal. Thus, the potential for speciation via adaptive radiation from a common ancestor through isolation, as witnessed in many of the terrestrial life forms (e.g. finches, tortoises, land snails and plants), is less likely to occur in the marine environment. Most marine populations exhibit a high level of genetic exchange within the Archipelago. This, however, is not to say that in-situ speciation has not played a role in shaping the unique diversity of marine life in Galapagos. Propagules arriving to Galapagos have become isolated over a sufficient period of time for character divergence to evolve via selection in response to different and/or changing environmental conditions. This process has resulted in rather high levels of endemism among many marine groups. For example, the proportion of endemism for well-studied invertebrate groups ranges from 17% in echinoderms to 38% in crustacean amphipods. These figures refer not only to derivations mainly from the Panama province but also forms whose closest relatives occur in the warm central and western Pacific region and along cool temperate South American shores (see Table 7.2). At the same time, some species share affinities with the Atlantic fauna, representing a divergence following the closure of the Central American Seaway about 3.1 million years ago (Keigwin 1982). This wide diversity reflects the fact that Galapagos lies in the path of major converging water masses. The frequency and abundance of propagules arriving via the various current systems would have occurred differently in past climatic eras. For example, when Galapagos was drier (cooler), or wetter (warmer), modulating environmental conditions would have increased the overall diversity of the fauna and influenced the evolution of island endemics. It should be noted, however, that unlike the terrestrial environment, the majority of unique biota found in


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Galapagos represents an evolutionary process of differentiation rather than one of diversification via adaptive radiation – as is more characteristic in the terrestrial environment. Hybridization The importance of hybridization to Galapagos marine biodiversity has been illustrated in recent studies of the demography and age structure of damselfish in the eastern Pacific (Arnold 1992). Collections made in Galapagos in 1995 revealed that a significant number of the common mainland dominant damselfish Stegastes acapulcoensis had invaded Galapagos during strong to very strong ENSO years and had hybridized with the native species Stegastes arcifrons (Meekan et al.1999, Meekan et al. 2001). Based on the number of yearly annuli on the earbones, these sample collections made in 1995 showed a large pulse of individuals arriving during strong and very strong El Niño events (1972-73; 1982-83; 1987 and 1990-1992). The collection also revealed one 32-year-old individual that had been recruited during the 1964 El Niño. While these species had come into contact on several occasions during recent times it was only in 1995 that hybridization was observed between these two species. The hybrids were identified as possessing heterozygous alleles at 4 loci. These studies indicate that immigration via invasion from the mainland to Galapagos can be driven by environmental perturbations, which are revealed in the population age structure of Stegastes acapulcoensis. While hybridization is generally thought to be detrimental to both species, natural hybridization may generate novel genotypes that could result in adaptive evolution and/or founding of a new evolutionary lineage. One could envision that some hybrids would fare better in a novel environment, such as might occur as a result of prolonged changes in climatic conditions. The extent that such events have molded past evolutionary trajectories of species in Galapagos is unknown. A thorough study of hybridization would certainly provide interesting insights into evolutionary processes that occur in insular environments. Extinction and colonization Recent ENSO warming anomalies, particularly the great 1982/83 El Niño, have had a profound effect on individual species, communities, and the ecosystem in Galapagos. What do the warmer waters of El Niño mean for ocean life? In Galapagos, starvation and death. The cold tongue of the Cromwell Current that moves along the equator during normal conditions upwells from the deep carrying nutrients into the surface waters where light can penetrate. Phytoplankton, the microscopic plants at the base of the marine food web, need both nutrients and light to grow. During ENSO conditions the warm surface waters act like a cap keeping the cold, nutrient-rich subsurface waters from reaching the euphotic zone. The lack of nutrients essentially starves the entire ecosystem beginning with the microscopic phytoplankton and extending all the way to the top of the food chain, marine mammals. One area particularly affected by El Niño is the Galapagos Archipelago. As a consequence of these warm-cold events, whole communities of marine plants and animals appear to have been lost from islands in Galapagos following the 1982/83 El Niño, and it is possible that some endemic marine species have now become extinct. No systematic studies have been conducted to document the decline and loss of such species, or to implement recovery plans for threatened marine plants and animals in the region. The devastation in Galapagos caused by the 1982/83 El Niño on overall marine biodiversity has been poorly documented but the scale is indicated by analysis of corals – the group of marine organisms best-studied locally. Glynn (1990, 1994) found that 97% of corals were lost from Galapagos during and immediately after the 1982/83 El Niño. Although individual colonies persist, the majority of corals were killed by extreme surface temperatures in 1982/83 (Glynn 1990). This mortality was followed by intense grazing by the sea urchin Eucidaris thouarsii, leading eventually to the total loss of reefs in the archipelago. Grazing activities in turn led to severe bioerosion and subsequent destruction of the coral carbonate structures which had been accreted by corals over the last several thousand years. Numerous reef-associated species presumably disappeared with the loss of coral reefs, although this has not been documented. Another negative consequence attributed to recent ENSO activity is the apparent extinction or near extinction of two planktonic-feeding fishes endemic to Galapagos and nearby oceanic islands: Azurina eupalama (Blackspot Chromis) and Acanthemblemaria castroi (Castro’s tube blenny). Azurina has not been seen since the 1982/83 ENSO. During that event the population of A. castroi declined in numbers but eventually recovered. Following the recent 1997/98 ENSO (1998) its populations were again decimated, however subsequent observations indicate that recovery is presently occurring.


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Another key example of loss of biodiversity associated with ENSO anomalies in Galapagos is that, prior to 1983, low intertidal shores in the central and southern islands were covered in dense beds of endemic brown algae belonging to the genera Sargassum and Bifurcaria. These conspicuous and formerly-dominant macroalgae, and most others amongst more than 100 seaweed species endemic to Galapagos, have not been recorded or seen in Galapagos waters in recent years. While the possibility remains that remnant populations still exist in the cooler western region around Fernandina and western Isabela, low intertidal algal communities have certainly disappeared from former algal strongholds in the east and south. The loss of endemic Galapagos seaweed species (and its species and communities) is of global significance, particularly as only one seaweed species worldwide is recognized to have become extinct (Vanvoustia bennettiana – an Australian species only recorded once from Sydney Harbour). The same is true for invertebrate marine species, many of which were formerly common but have not been seen since the 1983 and 1997 El Niños. For example, several intertidal species common in low shores of the south-central islands have disappeared or suffered massive population reductions. Among them, several low intertidal and shallow subtidal species, including the two endemic sunstars Heliaster cumingi (now an extremely rare species) and Heliater solaris (not seen since the 1982-83 El Niño). Included amongst possible extinctions are potentially-valuable aquaculture species, including the large (to 200 mm) endemic Magnificent Scallop species Nodipecten magnificus. This mollusk was formerly sufficiently common to be utilized for human consumption but only a few individuals have been seen during the past five years. For this large scallop species, the sudden growth of the dive fishery for sea cucumbers in the early 90’s may also have contributed to its rapid decline and local extinction. Enhanced immigration as well as extinction of species also apparently occurs during extreme ENSO anomalies. In recent times, several marine species from the central and western Pacific have been recorded recruiting to Galapagos after strong ENSO events. Following the 1982/83 El Niño, individuals of four species of western Pacific butterfly fishes were observed at several locations in the Archipelago: Chaetodon auriga (Threadfin Butterfly Fish), C. kleini (Klein’s Butterfly Fish), C. lunula (Raccoon Butterfly Fish) and C. meyeri (Meyer’s Butterfly Fish). Some persisted for a few years but apparently without successfully becoming established through local recruitment. Following the most recent ENSO of 1997/98, individuals of the Indo-Pacific wrasses, Thalassoma purpureum and Stethojulis bandanensis (Banded Wrasse) were observed. While T. purpureum (Surge Wrasse) was uncommon and largely remained confined to the northern islands, the banded wrasse enjoyed high levels of recruitment, reaching the status of common at several locations in the tropical eastern Pacific. Analyses of otolith increments indicated that a one-month long pelagic larval duration was sufficient for the species to spread among isolated island groups in this region. Adults of this species were present in Galapagos during the subsequent cold La Niña period. Shortly after the last ENSO 1997/98, several new fish arrivals were found: Encheliophis dubius (Pearl Fish commensal on Stichopus fuscus), Lutjanus colorado (Colorado Snapper), and Lutjanus guttatus (Snapper) (Bustamante and Rivera, pers. comm.). These findings, that range extensions occur during ENSO warm water conditions, and that adults can persist in their new locations, indicate that the barriers to successful colonization for some species in this region may hinge on specific critical aspects of the pelagic environment that are required by larvae. These critical aspects probably involve temperature, current flow, or other physical or biological conditions that may exist only during ENSO periods (Glynn and Ault, 2000). In addition to the above fish examples, recent colonization of invertebrates has also been recorded. Noticeable amongst these was the discovery of large individuals (>1 year old) of Panulirus albiflagellum (White Antenna Spiny Lobster) along the northern shores of Isabela after the 1997/98 ENSO. The invasion of this tropical species, normally found in New Caledonia and the western Pacific, indicates that the natural arrival of new species to the Galapagos marine environment is an active and dynamic evolutionary process (Bustamante et al. in prep.). The significance of these events in an evolutionary sense is that some of the species replacements and losses will shape communities and lead to evolutionary changes in species interactions. Dynamic species turnover witnessed on islands clearly provides important insights into how species evolve under changing conditions.


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While examples of immigration and extinction are expected to occur in isolated oceanic island environments, the scale of these changes, particularly recent levels of extinction, may be unique globally. We consider it likely that these large-scale changes to inshore marine communities are directly related to the fact that Galapagos lies near the epicenter of heating anomalies during ENSO events (Glynn & Ault 2000). Clearly, the relationship between extreme ENSO events and loss of marine biodiversity in Galapagos needs to be properly documented and further investigated as a matter of global importance. Such studies have direct relevance to predictions of loss of global biodiversity associated with the increasing frequency and magnitude of warming events associated with global climate change. REFERENCES

Abbott, D.P. 1966. Factors influencing the zoogeographic affinities of the Galpagos inshore marine fauna. The Galapagos. Proceedings of the Symposia of the Galapagos International Scientific Project (ed R. Bowman). Univ. Calif. Press, Los Angeles.

Allan, R., J. Lindesey & D. Parker 1996. El Niño Southern Oscillation and climatic variability. CSIRO, Canberra.

Arnold, M.L. 1992. Natural hybridization as an evolutionary process. Ann. Rev. Ecol. Syst. 23:237-261.

Bustamante, R.H., G.M. Branch, S. Eekhout, B. Robertson, P. Zoutendyk, M. Schleyer, A. Dye, N. Hanekom, D. Keats, M. Jurd & C. MacQuaid 1995. Gradients of Intertidal Productivity Around the Coast of South Africa, and their Relationship to Consumer Biomass. Oecologia 120: 189-201

Cairns, S.D. 1986. Stylasteridae (Hydrozoa: Hydroida) of the Galapagos Islands. Smithsonian Contrib. Zool. 426: 42 pp.., 27 figs.

Chavez, F.P. & R. C. Brusca 1991. The Galapagos Islands and their relation to oceanographic processes in the tropical Pacific. In Galapagos Marine Invertebrates ( ed. M.J. James) pp. 9-33. Plenum Press, N.Y.

Christie, D.M., R.A. Duncan, A.R. McBirney, M.A. Richards, W.M. White, K.S. Harpp & C.G. Fox 1992. Drowned islands downstream from the Galapagos hot spot imply extended speciation times. Nature 355:246-248.

Colinvaux, P.A. 1972. Climate and the Galapagos Islands. Nature 240:17-20.

Cushing, D.H. 1989. A difference between ecosystems in strongly stratified waters and in those that are only weakly stratified. J. Plankt. Res . 11:1-13.

Dunbar, R.B., G.M. Wellington, M.W. Colgan & P.W. Glynn. 1994. Eastern Pacific sea surface temperatures since 1600 A.D.: The delta 18O record of climate variability in Galapagos corals: Paleoceanography, v. 9, pp. 291-315.

Emanuel, B. P., R. H. Bustamante, G. M. Branch, S. Eekhout & F. J. Odendaal 1992. A zoogeographic and functional approach to the selection of marine reserves on the west coast of South Africa. S. Afr. J. Mar. Sci. 17:1017-1030.

Farnsworth, E.J. & A. Ellison. 1997. The global status of mangroves. Ambio 26: 328-334.

Garth, J. S. 1991. Taxonomy, distrubution, and ecology of Galapagos Brachyura. In Galapagos Marine Invertebrates – Taxonomy, Biogeography and Evolution in Darwin’s Islands (ed. James, M.J.) pp. 123-142. Plenum Press, New York.

Genin. A. P.K. Dayton, P.F. Lonsdale, & F.N. Spiess 1986. Corals on seamounts peaks provide evidence of current acceleration over deep-sea topography. Nature 322: 59-61.


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Glynn, P.W. 1990. Coral mortality and disturbances to coral reefs in the tropical Eastern Pacific. In Global ecological consequences of the 1982-83 El-Niño-Southern Oscillation (ed. P.W. Glynn). Elsevier Press, Amsterdam.

Glynn, P.W. 1994. State of Coral reefs in the Galapagos Islands: natural vs anthropogenic impacts. Mar. Bull. Poll. 29:131-140.

Glynn, P.W., G.M. Wellington, & C. Birkeland 1979. Coral reefs growth in the Galapagos: Limitations by sea urchin. Science 203: 47-49.

Glynn, P.W. & G. Wellington 1983. Corals and coral reefs of the Galapagos Islands. University of California Press, Berkeley.

Glynn, P.W. & J.S. Ault. 2000. A biogeographic analysis and review of the far eastern Pacific coral reef region. Coral Reefs 19:1-23.

Harris, M.P. 1969. Breeding season of sea-birds in the Galapagos Islands. J. Zool. (Lond), 159: 145-165.

Houvenaghel, G. T. 1984. Oceanographic Setting of the Galápagos Islands. In Key Environments: Galápagos (ed. R. Perry). Pergamon Press, Oxford.

INOCAR, 2000. Tablas de mareas y datos astronómicos del sol y la luna. Instituto de Oceanografico de la Armada. Guayaquil.

James M. J. (ed.) 1991. Galapagos Marine Invertebrates – Taxonomy, Biogeography and Evolution in Darwin’s Islands. Plenum Press, New York.

Jennings, S., A.S. Brierley & J.W. Walker 1994. The inshore fish assemblages of the Galapagos Archipelago. Biol. Conserv. 70:49-57.

Kay, E.A. 1991. The marine mollusks of the Galapagos, Determinants of insular marine faunas. In Galapagos Marine Invertebrates – Taxonomy, Biogeography and Evolution in Darwin’s Islands (ed. M.J. James) pp. 235-252. Plenum Press, New York.

Keigwin, L.D.,Jr. 1982. Isotopic paleoceanography of the Caribbean and east Pacific: Role of Panama uplift Late Neogene time. Science 217:350-52.

Longhurst, A.R. 1985. Relationship between diversity and the vertical structure of the upper ocean. Deep Sea Res. 32: 1535-1570.

Lonsdale, P. 1977. Clustering of suspension-feeding macrobenthos near abyssal hydrothermal vents at the oceanic spreading centers. Deep Sea Res. 24: 857-863.

MacArthur, R.H. & E.O. Wilson 1967. The theory of island biogeography. Princeton, Princeton University Press.

McCosker, J.E., R.H. Bustamante and G.M. Wellington. In prep. The Freshwater Eel, Anguilla marmorata, discovered at Galápagos (submitted to Noticias de Galápagos).

McCosker, J.E. & R.H. Rosenblatt 1984 Marine Environment and Protection. In Key Environments, Galapagos (ed. R. Perry ) pp. 133-144. Pergamon Press.

Meekan, M.G., G.M. Wellington & L. Axe 1999 El Nino-Southern Oscillation events produce checks in the otoliths of coral reefs in the Galapagos Archipelago. Bull. Mar. Sci. 64:383-390.

Meekan, M.G., J.L. Ackerman & G.M. Wellington 2001. Demography and age structures of coral reef damselfishes in the topical eastern Pacific. Mar. Ecol. Prog. Ser. 211:223-232.

Monniot, C. & F. Monniot. 1989. Ascidians collected around the Galapagos by the Johnson Sea-Link Research submersible. Proc. Biol. Soc.. Wash. 102: 14-32.


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Silva, P.C. 1964. Status of our knowledge of the Galapagos benthic marine algal flora prior to the Galapagos International Scientific Project. In The Galapagos Islands Symposium of the Galapagos International Scientific Project (ed. R.I. Bowman) pp. 149-156. University of California Press, Berkeley.

Snell, H.M., P.A. Stone & H.L. Snell 1995. Geographical characteristics of the Galapagos Islands. Noticias Galapagos 55: 18-24.

Wellington, G.M. 1984. Marine Environment and Protection. In Key Environments. Galapagos (ed. R. Perry), pp. 247-263. Pergamon Press.

Wellington, G. M., A. E. Strong & G. Merlen 2000. Sea surface temperature variation in the Galapagos Archipelago: a comparison between AVHRR nighttime satellite data and in-situ instrumentation (1982-1988). Bull. Mar. Res. in press.

Witman, J.D. & F. Smith Rapid community change at a site in the Galapagos Marine Reserve. In press, Biodiversity and Conservation.

CDF/WWF Biodiversity Vision Chapter 8 - Marine Conservation Criteria

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CHAPTER 8 – CONSERVATION CRITERIA FOR THE MARINE BIOME

Principal Authors - G.M. Branch, J.D. Witman, R. Bensted-Smith, R.H. Bustamante, G.M. Wellington, F. Smith & G.J. Edgar.

At the workshop the marine group defined three principal criteria for marine conservation: (1) the total protection of representative areas covering all biogeographic provinces in the region and all habitats present within them, (2) the maintenance of the composition and functioning of the ecosystem, and (3) the value and sustainability of products and services of the ecosystem. These reflect the need for conservation of biological communities, of ecological processes, and of species, respectively.

CRITERION 1: Extent to which biogeographic zones and habitats are represented in No-Take-Areas (NTAs) and surrounded by areas with effective management.

If all biogeographic regions and all habitats within them receive adequate protection, this will ensure that the majority of species and the functioning of ecosystems will be conserved. This approach serves as a surrogate for identifying 'hot-spots' where species richness and endemism are high, and is more effective in ensuring a representative coverage of all systems (Hockey and Branch 1997; Roberts et al. in press a,b). In defining indicators for this criterion, it was decided to treat deep, offshore waters separately from the coastal waters and other shallow areas. Most of the evidence for distinct biogeographic zones has been supported by data on coastal biodiversity and very little is known about open ocean biodiversity distribution patterns (Ekman, 1953; Pielou, 1979, 1983; Vermeij, 1978, 1992; Murray and Litter, 1981; Gosliner, 1987; Joosten and Van der Hoek, 1986; Kohn, 1990). In the case of the Galapagos Archipelago, congruent biogeographic divisions have been demonstrated for sea birds, reef fish and benthic invertebrates and seaweed (Harris, 1969; Jennings and Brierly, 1994; Bustamante et al. unpubl. data). However, there is reported evidence in which large-scale biogeographic units based on coastal biota, are also “replicated” or confirmed by oceanic and planktonic communities (Barange et al. 1992, Gibbons et al. 1995). Furthermore, in the case of Galapagos, a recent study (Banks, 1999) has demonstrated a close correlation between the surface temperature of Galapagos water masses and the biogeographic divisions proposed by Harris (1969). Thus, it was decided that for the evaluation and assessment of Criterion 1, the current knowledge strongly confirms the existence of 3-4 biogeographic units. Further work is needed to define boundaries and numbers, but the model proposed by Harris (1969) presently appears valid and is used for the assessment of this criterion. In the Galapagos, coupled pelagic and open ocean biota are expected to follow similar abundance and distribution patterns.

With this in mind, the indicators for Criterion 1 are:

• The proportion of distinct habitats in each biogeographic zone, which have >20% of their area in no-take areas (NTAs). This refers to all habitats, both coastal and open-ocean, in each biogeographic zone.

• The proportion of the total area of all near-shore and other key habitats, principally the shallow sea mounts (“bajos”), which lie within effectively protected NTAs.

• The proportion of NTAs, which are surrounded by well-regulated fisheries and sound environmental management.

The concepts underlying Criterion 1 have not been thoroughly tested and used so far for marine conservation. Instead, marine biodiversity protection and conservation has been based largely upon traditional fisheries management criteria, such as quotas, effort control, size, season etc., all of which are aimed at individual species, not entire habitats or ecosystems. These management tools have failed consistently worldwide (FAO, 1995; Lubchenco et. al. 1995; Roberts, 1997; Vitousek et. al. 1997; Pauly et. al. 1998). This situation has led managers and scientists to initiate efforts to develop new and innovative ways to conserve and manage


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exploited marine resources (Fujita, 1998). Marine NTAs are being established worldwide in response to growing recognition of the conservation crisis that is now evident in the oceans. Their value has yet to be demonstrated on a large scale, but the examples so far (of which there are over a hundred) have consistently shown the recovery and enhancement of species and communities (Halpern et. al. 2000). With this evidence, scientists are proposing that the creation of substantial NTAs on a worldwide scale will provide a way to achieve simultaneously the apparently conflicting goals of effective protection and sustainable utilization of marine resources. Recent advances in this area of marine research and conservation, have shown that networks of NTAs represent a viable solution for sustaining fishery populations and marine ecosystems (Davis et. al., 1991; Russ and Alcala, 1998a,b; Roberts, 1995; Ballantine, 1997; Castilla and Fernandez, 1998; Murray, et. al. 1999; Nowlis and Roberts, 1997). Despite this, NTAs have only in a few cases been tested on a regional scale, where complex biogeographic units may exist (see Attwood et al. 1997a,b). In most parts of the world, multiple divisions or breaks occur between biogeographic provinces, and hundreds or even thousands of kilometres may separate units (Ekman, 1953; Murray and Littler, 1981; Roberts, et. al. 1992; Vermeij, 1992; Emanuel, et al. 1993; Barry, et. al. 1995, Bustamante & Branch 1996). However, in the case of Galapagos, a unique opportunity exists to assess the application and value of this criterion for biodiversity protection, due to the archipelago’s relatively small geographical scale, with biogeographic units just tens of kilometers apart.


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CRITERION 2 – Extent to which composition and functioning of the ecosystem are maintained.

Another scientific criterion, which will serve as a benchmark for the protection of the Galapagos marine ecosystem, is the extent to which the composition, functioning and services of the ecosystem are maintained. This approach recognizes that (1) each ecosystem is composed of individual organisms, populations, and communities that interact with each other and the physical environment; and (2) there are ecological relationships or processes, such as trophic (feeding) relationships, competitive interactions, positive (mutualistic) interactions, and source-sink (recruitment) relationships. All these aspects characterize the functioning of the Galapagos marine ecosystem. These ecological functions greatly influence contemporary levels of biodiversity in the Galapagos and are necessary considerations.

Many ecological relationships are dependent ones where the survival of a population in a given marine habitat depends on: (i) the supply of prey or nutrients, (ii) the supply of larvae or reproductive propagules, and (iii) the persistence of a species to continue mutualisms. In this sense, marine communities in the Galapagos are strongly linked by dependent relationships. Thus, widespread changes can occur throughout the ecosystems if important species involved in these relationships are removed or drastically reduced in abundance. This idea is illustrated below, in Fig. 8.1.

Top Figure 8.1. Diagram illustrating a simplified

version of a Galapagos marine food web with orcas (O) and sharks (S) depicted as top predators:

4th. sea lions (s), piscivorous fish (P) such as

groupers, jacks and barracuda, predatory birds (B = cormorants, pelicans, boobies) as fourth level consumers;

3rd lobsters (L), crabs (C), gastropods (snails, G),

and herring and anchovies (H) as tertiary consumers;

2nd. sea urchins (U), herbivorous fish (F)

herbivorous mollusks (M), barnacles (b), iguanas (I) as secondary consumers;

1st. zooplankton (z) as primary consumers; Primary Producers macroalgae (A) and

phytoplankton (P) as primary producers. The grazing trophic level is comprised of primary and secondary consumers.

C

s

L

B

M

G

A

b F

Z

P

U

P

I

H

S

O


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The original food web (above) has been altered by nearly eliminating herbivorous fish (surgeonfish, parrotfish and damselfish) to show how changes in trophic relationships can have widespread effects on the Galapagos ecosystem. The size of the circles is indicative of the abundance of the feeding group. Note that the abundance of species in the upper two trophic levels has decreased and that the abundance of algae (brown, green and red seaweed and algal turf) has increased due to the reduction of herbivorous fish. The components of the food web that changed are shaded in gray.

Such changes are hypothetical in this Galapagos marine example, but they have occurred in Caribbean reef ecosystems where over-fishing of herbivorous fish has contributed in part to a large increase in the abundance of benthic algae (Hughes 1994). Macroalgae have replaced coral as the most abundant species at some depths on Jamaican reefs (Andres and Witman 1995).

The principal indicator for measuring the status of ecological composition and function, i.e. Criterion 2, is as follows:

• The status of a selected taxa chosen to cover a range of the following variables:

- representation of trophic levels; - potential as keystone species (i.e. those with critical ecological functions); - attributes of their life history, such as the distances of larval dispersal, which influences

the degree of connectivity among populations at different sites; - sensitivity to environmental change (including both sensitive and insensitive species); - whether they are targets of exploitation (including both exploited and unexploited

species); - endemism or rarity; - importance for connecting marine and terrestrial ecosystems.

In the workshop the marine group identified on this basis a group of taxa that together could serve as indicators. A preliminary table of taxa appears in Annex 8.1. The priorities are:

• Seaweed • Corals (hermatypic) • Black corals / Gorgonians • Sea urchins (Tripneustes, Eucidaris) • Marine Iguanas • Grazing fish (parrot fish, surgeon fish and damsel fish) • Predatory fish: groupers, snappers and jacks (including Mycteroperca olfax) • Lobsters • Sea cucumbers • Sharks (Hammerhead and others) • Barnacles / Ascidians • Predatory snails / Conch • Cetaceans • Pinnipeds (fur seals and sea lions)

Co

L

M

G

b

Z

U

P

I

H

A


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Additional indicators for Criterion 2 are:

• The total diversity and abundance of species within each biological community and conversely, known cases of extinction.

• Genetic diversity within small populations at risk of extinction (low genetic diversity can increase susceptibility to extinction - Soulé 1980).

• Presence, abundance and distribution of alien species and their impact on native species • Prevalence and distribution of pathogens and parasites. • Resilience of biological communities and species, which is measured by the rate at which

they recover from disturbance or depletion. • Connectivity between sites, and local population replenishment within sites, measured by

the levels of recruitment of selected species. • Extent of anthropogenic habitat modification (including changes in structure, composition,

water quality etc.). • Maintenance of land-sea interactions

CRITERION 3 – The value and sustainability of products and services.

The marine ecosystem, with its composition and function intact, provides important services for the local community and humanity at large: economic utilization (extractive or non-extractive), food, clean water, sediment retention, detoxification and sequestration of pollutants and recycling of nutrients and other such services (Daily, 1997).

Many of these services operate in the Galapagos marine ecosystem over a local scale, for example: protection of coastlines by mangroves and coral reefs from the battering of waves; trapping and filtration of sediments provided by estuarine plants (reed beds, salt marshes and mangroves); filtration of water in bays by filter feeders such as sponges, ascidians and bivalves; creation of critical habitats required for spawning, recruitment or growth of biota, for example mangrove roots; maintenance of habitats for juvenile and adult stages of biota; and the provision of places desirable for recreation, inspiration, education and the advancement of knowledge. Other services of a thriving marine ecosystem in Galapagos operate over a regional scale, for example: generation and maintenance of biodiversity that is fundamental to tourism; decomposition of dead biological material; and recycling of nutrients.

Therefore, the third criterion for assessing conservation of Galapagos marine biodiversity refers to the value and sustainability of these benefits and has the following set of indicators. The indicators are focused mainly on the status of economically valuable species. Perhaps indicators for other services, such as the provision of habitat for juvenile fish or recycling nutrients, should be added at a later stage.

• Abundance and population structure of economically important species, in particular large marine animals, seabirds, reef fish and invertebrates.

• Number of trophic levels and mean trophic level of exploited species. • Catch Per Unit Effort (CPUE) of exploited species. • Spawning stock, measured as biomass per recruit. • Levels of the indirect impacts of fishing, tourism, waste disposal or other activities such

as by-catch, pollution, and physical damage to corals. • Level of benefit relative to volume of extraction or frequency and intensity of non-

extractive uses. • Level and orientation of subsidies (“orientation” refers to whether the subsidies are

positive, i.e. they encourage conservation, or perverse, i.e. they foster overfishing).

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Attwood, C.G., Mann, B.Q., Beaumant, J. and J.M Harris. 1997b. Review of the state of marine protected areas in South Africa. S. Afr. J. mar. Sci. 18: 341-367.

Ballantine, W.J. 1997. Design principles for systems of ‘no-take’ marine reserves. Paper for workshop: The Design and Monitoring of Marine Reserves at Fisheries Center, University of British Columbia, Vancouver, Feb 1997.

Banks, S.A. 1999. The use of remotely sensed AVHRR data in determining SST variability and zonation across the Galapagos Marine Reserve. M.Sc. thesis in Oceanography at Southampton Oceanography Centre. 46 pp.

Barange, M. Pillar, S.C. and L. Hutchings. 1992. Major pelagic borders of the Benguela upwelling system according to euphausiid species distribution. S. Afr. J. mar. Sci. 12: 3-17.

Barry, JP; Baxter, CH; Sagarin, RD; Gilman, SE. 1995. Climate-related, long-term faunal changes in a California rocky intertidal community. Science 267 (51):672-675.

Bustamante, R. H. and G. M. Branch 1996. Large-scale patterns and trophic structure of Southern African rocky shores: the roles of geographic variation and wave action. Journal of Biogeography, 23:339-351.

Castilla, J.C. and M. Fernandez. 1998. Small-scale benthic fisheries in Chile: on comanagement and sustainable use of benthic invertebrates. Ecological Applications 8: S124-S132.

Daily, G. C. 1997. Nature’s services, societal dependence on natural ecosystems. Island Press, Washington D.C., USA.

Davis, Gary E., Jameson, S. C. and J. E. Dugan. 1991. “Potential Benefits of Harvest Refugia in Channel Islands National Park and Channel Islands National Marine Sanctuary”. pp. 2962-2972 in: O. Magoon et al (eds.) Proceedings of the Seventh Symposium on Coastal and Ocean Management. American Society of Civil Engineers, New York, NY.

Ekman, S. 1953. Zoogeography of the sea. Sidwick and Jackson limited, London, United Kingdom.

Emanuel, B. P., R. H. Bustamante, G. M. Branch, S. Eekhout and F. J. Odendaal, 1992. A zoogeographic and functional approach to the selection of marine reserves on the west coast of South Africa. S. Afr. J. mar Sci. 17: 1017-1030.

FAO. 1995. The state of the world's fisheries and aquaculture. FAO Fisheries Department. Food and Agriculture Organization of the United Nations, Rome.

FAO. 2000. The state of world fisheries and aquaculture in 2000. Food and Agriculture Organization of the United Nations http://www.fao.org/docrep/003/x8002e/x8002e00.htm (2000).

Fujita, R.M., T. Foran, and I. Zevos. 1998, Innovative approaches for fostering conservation in marine fisheries. Ecological Applications 8: S139-S150.

Gibbons, M., Barange, M., and L. Hutchings. 1995. Zoogeography and diversity of euphausiids around southern Africa. Mar. Biol. 123: 257-268.

Harris, M.P. 1969. Breeding season of sea-birds in the Galapagos Islands. J. Zoology (Lond), 159: 145-165.


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Hockey, P.A.R. and G.M Branch. 1997. Criteria, objectives and methodology for evaluating marine protected areas in South Africa. S. Afr. J. mar. Sci. 18: 369-383.

Hughes, T.P. 1994, Catastrophes, phase shifts and large scale degradation of a Caribbean coral reef. Science 265: 1547-1551.

Lubchenco, J., G.W. Allison, S.A. Navarrete, B.A. Menge, J.C. Castilla, O. Defeo, C. Folke, O. Kussakin, T. Norton, and A.M. Wood. 1995. Biodiversity and ecosystem functioning: Coastal systems. Pages 370-381 In: Global Biodiversity Assessment, Cambridge University Press, Cambridge, UK.

Murray, S. N., and M. M. Littler. 1981. Biogeographical analysis of intertidal macrophyte floras of southern California. Journal of Biogeography 8: 339-351.

Murray, S. N., R. F. Ambrose, J. A. Bohnsack, L. W. Botsford, M. H. Carr, G. E. Davis, P. K. Dayton, D. Gotshall, D. R. Gunderson, M. A. Hixon, J. Lubchenco, M. Mangel, A. MacCall, D. A. McArdle, J. C. Ogden, J. Roughgarden, R. M. Starr, M. J. Tegner, M. M. Yoklavich. 1999. No-take Reserve Networks: Sustaining Fishery Populations and Marine Ecosystems. Fisheries 24(11): 11-25.

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Pielou, E. C. 1983. Spatial and temporal change in biogeography: gradual or abrupt? Pages 29-55 in R. W. Sims, J. H. Price and P. E. S. Whalley editors. Evolution, time and space: the emergence of the biosphere. Academic Press, London, United Kingdom.

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Roberts, C.M. 1997. “Connectivity and Management of Caribbean Coral Reefs”. Science. Vol. 278: 1454-1457.

Roberts, C.M., S. Andelman, G.M. Branch, R.H. Bustamante, J.C. Castilla, J. Dugan, B. Halpern, K.D. Lafferty, H. Leslie, J. Lubchenco, D. McArdle, H. Possingham, M. Ruckleshaus, and R. Warner In press a. Ecological criteria for evaluating candidate sites for marine reserves. Ecological Applications.

Roberts, C.M., G.M. Branch, R.H. Bustamante, J.C. Castilla, J. Dugan, B. Halpern, K.D. Lafferty, H. Leslie, J. Lubchenco, D. McArdle, M. Ruckleshaus, and R. Warner In press b. Application of ecological criteria in selecting marine reserves and developing reserve networks. Ecological Applications.

Russ, G.R. and Alcala, A.C. 1998. Natural fishing experiments in marine reserves 1983-1993: community and trophic responses. Coral Reefs 17, 383-398.

Russ, G.R. and Alcala, A.C. 1998. Natural fishing experiments in marine reserves 1983-1993: roles of life history and fishing intensity in family responses. Coral Reefs 17, 399-417.


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CDF/WWF Biodiversity Vision Chapter 9 – Marine Biodiversity Status

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CHAPTER 9 – THE STATUS OF AND THREATS TO MARINE BIODIVERSITY

Principal Authors – R.H. Bustamante, G.M. Branch, R. Bensted-Smith and G.J. Edgar.

9.1 Constraints on Defining the Pristine State of Galapagos Marine Ecosystems

In the fluctuating marine envi ronment of Galapagos, which has been intensively exploited for whales and seals in the past century and increasingly during the past century for seafood, it is difficult to establish a reference point for a pristine state of marine biodiversity. However, we know that for most of the recently exploited species the intensity of extraction has increased substantially in the past decade or two, and some of the changes caused by that extraction have been recorded. There are also some species whose depletion by hunting (whales and fur seals) or by predation by cats and rats on land (dark-rumped petrels, marine iguanas) occurred earlier in the Islands’ human history. Thus, it is possible to provide a description of current marine biodiversity, and to identify biodiversity changes known to have been caused by early visitors or, more recently, by El Niño, fishing or other factors. Even this approach is limited by the fact that the marine flora and fauna, especially those in deep water, have been little studied compared to their terrestrial counterparts.

Perhaps the most important point to bear in mind, when considering what the pristine state of Galapagos may have been like, is the dynamic nature of the marine ecosystem and its highly variable environment, as described in Chapter 7. Thus any anthropogenic trends are superimposed on significant natural fluctuations.

9.2 The Current Status of Galapagos Marine Biodiversity

This section assesses the status at the time of the workshop in May 1999, with some updated information added from 2000 and 2001.

Like most of the world’s marine ecosystems, the seas of the Galapagos Islands have felt the cumulative impacts of human activities for several centuries. The marine ecosystems contained in coastal, shallow and deeper waters, have received the impacts of hunting, fishing, coastal development, pollution and tourism. Consequently, the ecosystems are by no means in a pristine state, however that may be defined. Nevertheless, the low resident human population and the relatively small scale of most impacts (with a few exceptions, such as whaling), have allowed Galapagos to retain most of its marine biological diversity. However recent surveys by CDRS have revealed major changes in biological communities and severe declines of some marine species, with possible local – or even total- extinction in some cases. These changes appear to be a consequence of the exceptional severity of the 1983 and, to a lesser extent, the 1997/98 El Niño events, perhaps interacting with anthropogenic factors such as fishing (Edgar et al. in prep.).

El Niño events are a global phenomenon, the effects of which are observed most frequently in the region of the Eastern Pacific bounded by the Galápagos Islands, Cocos Island and the Ecuador coast. The focus of intensity is centered on Galápagos, with, for example, water temperatures off Puerto Ayora, Galápagos not declining below 25oC over an 18-month period during the 1997/98 El Niño, in comparison with water temperatures falling below 20oC during the winter months of normal years (CDRS water temperature monitoring data). In addition to greatly elevating water temperature and virtually eliminating its regional variation, El Niños also cause pronounced stratification of oceanic waters. They prevent nutrient upwellings, resulting in rapid declines in the availability of plant nutrients, which in turn leads to greatly diminished phytoplankton productivity and a catastrophic diminution in the base of the marine food chain. Most marine species then need to migrate, if sufficiently mobile, or cope with food shortage and starvation.

The impacts of recent El Niño events in Galápagos are poorly known for most groups of marine plants and animals, but some endemic species, which declined sharply during the extreme 1983 and 1997/98 El Niño events, have shown few signs of subsequent population recovery.


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Comparisons of baseline data presented in Wellington (1975) with data obtained in recent CDRS surveys show many differences. Wellington described beds of fucoid algae as dominant in inter-tidal and shallow habitats on the southern shores of most islands and the western shores of the westernmost islands of Isabela and Fernandina. He recorded densities of the sea urchin Eucidaris thouarsii at up to 15 m-2, and noted that barnacles “don’t compete well with more abundant algae”. By contrast, recent shallow-water surveys indicate a predominance of coralline-algal-encrusted rock dominated by sea urchins or barnacles, with localised patches of cropped turf algae, a five-fold increase in densities of E. thouarsii, and no fucoid algae at any of the >250 sites investigated using quantitative transects.

The recent CDRS surveys also failed to detect a number of endemic species considered common prior to 1983. For example, prior to 1983 many shores were covered by dense beds of endemic brown seaweeds (Bifurcaria galapagensis and several Sargassum species) but this formerly dominant macroalgal species (and others amongst more than 100 seaweed species endemic to Galápagos) has not been recorded anywhere in the Galápagos since the 1980s. It is likely that the loss of these seaweed beds has been accompanied by the decline or loss of associated communities of grazing invertebrates. Whereas declines in terrestrial plant communities are generally quickly noticed and threats identified, wholesale changes in marine plant communities appear to have taken place in Galapagos without documentation.

Similarly, a number of marine invertebrate and fish species, some of which were formerly common, have not been recorded during recent surveys, with some not having been sighted since the 1983 El Niño. Amongst these species are the endemic intertidal seastar Heliaster solaris and the blackspot chromis Azurina eupalama, a plankton-feeding damselfish (Roberts and Hawkins, 1999). Pronounced declines in populations of coral species were also detected during the studies of Glynn (1990, 1994), who found that 97% of corals were lost from Galapagos during and immediately after the 1983 El Niño and that little recovery has since occurred.

Salazar (2002) estimated in 2001 the total populations of sea lions (Zalophus wollebaeki) and fur seals (Arctocephalus galapagoensis) in Galapagos to be 16,000 and 6,000 individuals respectively and suggests that there has been a significant decline in comparison to the 1977-78 surveys of Trillmich, who estimated the total populations to be 40,000 and 30-40,000 respectively. Some locations, which in 1977/8 had large numbers of animals, were found in 2001 to be reduced to a few individuals. The methodologies used in the two surveys differed significantly and there are potentially large errors in the estimates of total populations, so great caution must be exercised in quantifying the population changes. Nevertheless, the available data does suggest a decline in both species, and the estimated total populations, especially in the case of the fur seal, are low enough to warrant urgent research to verify population status and trends, together with precautionary conservation measures.

A critical consideration in identifying the causes of the changes in biodiversity described above is the potential interaction between the ecological effects of El Niño and human activities, such as fishing and the introduction of invasive species. The last two decades, during which considerable changes in biodiversity have occurred, have also seen major growth in the human population of Galapagos and, in particular, in fishing, including both coastal artisanal fishing and offshore industrial fishing. It remains unclear whether the stresses on animal populations produced by extreme warming events are additive or multiplicative when compounded with fishing and other human-induced stresses, such as the presence of introduced species in coastal breeding areas. For example, a natural tendency is for fishers to increase effort to maintain income levels during El Niño years of low fish productivity; however, the marine ecosystem is presumably least able to cope with additional stresses at this time. Similarly, the relative contributions of fishing and climate change to recent changes in ecosystem structure remain unclear. For example, currently available information is insufficient to determine how much increasing sea urchin populations have been caused by climatic variations and how much by the functional removal of urchin predators such as lobsters and grouper.


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9.3 Threats to Marine Biodiversity and Opportunities for Conservation

Opportunities

The main opportunities for improving marine biodiversity conservation are: (1) the legal framework, the main goals of which are biodiversity conservation and sustainable utilization of resources; (2) the ongoing establishment of regulations and mechanisms to limit access to the fisheries, to allow only artisanal fishing, and to set fishing effort at sustainable levels for the long-term benefit of the fisheries; (3) the legal requirement and local agreement to implement a network of No-Take-Areas (NTAs) and to zone the remaining areas for specific activities and types of control; and (4) an institutionalized participatory process for scientifically-based conservation and management of Galapagos’s marine biodiversity, with access limited to defined user groups.

The legal framework established by the Ecuadorian Government in the past two years for the Galapagos Marine Reserve (GMR) provides a unique opportunity to implement new and innovative conservation and management measures for conserving marine biodiversity. As mentioned in Chapter 2, the Special Law for Galapagos (SLG) of 1998 created the GMR, within which the only extractive activity is local artisanal fishing. The law curbs migration by restricting citizen rights for residence in the islands, in order to protect the biodiversity and economic base of the islands (mainly tourism and fishing). Having thus restricted the groups of people who may have access to marine resources, the law institutes a mechanism whereby local stakeholders (fishing sector, tourism sector, scientists, conservationists and environmental educators) can participate in the planning, management and decision-making of the Reserve, including the elaboration of the management plan (approved in March 1999).

The participatory management process has had many ups and downs but has achieved a great deal and has proved resilient since it was set up as an informal grouping in 1997. Indeed, the creation of the Marine Reserve under the management of the Galapagos National Park Service, with boundaries extending 40 miles from the archipelago and with industrial fishing prohibited, is due in large measure to the CDRS-GNPS consensus-building initiative, that was the precursor to the participatory management regime. Already the local “Participatory Management Board” routinely uses fisheries monitoring and survey data, much of it collected together with stakeholders, in the negotiation of fisheries seasons, quotas and rules. The provisional zonation of the GMR (see below), though not ideal, has the great merit of having been decided by consensus. A recent evaluation by IUCN confirmed the professionalism and innovatory nature of the participatory management system for the Galapagos Marine Reserve (G. Borrini-Feyerabend and M. Taghi Farvar, pers. comm.).

To this opportunity provided by law and by the institutionalised participatory management system, may be added some other advantages. Compared to fishing communities associated with most large marine protected areas, Galapagos fishers are few in number, which facilitates regulation, communication and educational activities. They are distributed between just four towns and are prohibited from landing onshore around at least 90% of the coastline. The Galapagos National Park Service, having been given in 1998 responsibility for the administration and management of the GMR, has steadily built up its enforcement and management capability, whilst cooperation with the Ecuadorian Navy has been increasing during 2001. The Government of Ecuador is taking on an Inter-American Development Bank loan, US$ 5 million of which is earmarked for the GMR. In December 2001 the Government’s proposal to have the Galapagos World Heritage Site extended to include the GMR was accepted by the World Heritage Committee.

The central management tool of the management plan, to achieve Galapagos marine conservation, is the approved provisional zoning scheme (Figure 9.1). This scheme was designed to protect directly threatened marine biodiversity, but also to reduce conflicts between uses, principally tourism, fishing and scientific research. This provisional zoning scheme will be evaluated and modified after two years of full implementation (start date is unclear but is probably in 2002, as physical demarcation has been delayed) and then again after four years, after which it becomes a permanent conservation and management provision. The zoning scheme for coastal and nearshore areas contemplates basically four main types of zones:

• strict no-take and no-go zones, where only scientific use is allowed (2.1);


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• no-take zones where tourism, recreation and education are permitted (2.2); • extraction zones under traditional fisheries management regimes but with the possibility of

more restrictions than occur in deeper waters (2.3); and • areas with rotating closures or under special management (2.4).

The NTAs include a range of replicated sites of different sizes and distances apart, which form a network representing most habitats in each biogeographic zone. Thus, the zoning scheme will implement a network of zones with take and no-take regimes that simultaneously serve multiple goals, i.e. to achieve protection and allow sustainable resource utilization. The diagram in Figure 9.1 depicts the provisional agreement on zoning, achieved in April 2000. Some details of particular sites under zone 2.2 and the detailed planning of zones 2.4 remain to be worked out. Just over 17% of the coastline is designated as No Take Areas (2.1 or 2.2), with some small additional NTAs to be designated in the subsequent mini-zonation of the areas near ports (2.4). Currently, no NTAs have been delineated in deep-water habitat more than 2 miles offshore.

Figure 9.1. General map of the provisional coastal zoning scheme for the GMR approved in April 2000. The zones are; 2.1 fully-protected ‘no-take’ area, in green; 2.2 non-extractive use areas, in blue; 2.3 regulated extractive uses, in red; and 2.4 special zones nearby the inhabited port areas, in black. The percentages for each zone will change slightly following definition of the exact location and extent of zones 2.2 and 2.4.

Threats

Fishing

The principal anthropogenic threat to Galapagos marine biodiversity is inappropriate and/or excessive fishing. This is no surprise: around the world, intensive and permanent fishing at small and large scales, is widely considered to be the most important agent of ecological change in the sea. Over the years, fishing has systematically stripped the highest trophic levels from marine food webs, eliminating some of the most valuable and spectacular of the marine fauna. Intense or persistent fishing on marine food webs has been shown to disrupt the composition and functioning of marine communities and ecosystems (Pauly and Christensen, 1999, Jackson et al. 2001), from kelp forests (Dayton et al. 1998) to coral reefs (Roberts 1995). In Galapagos, as already discussed, fishing pressure is super-imposed on the effects of the ENSO cycle, which may be intensifying. The consequences for endemic marine life of the combination of effects appear extremely serious.


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The fact that Galapagos fisheries have hitherto operated on a relatively small scale has not prevented the over-exploitation of target resources. Recent modelling studies have shown that the current artisanal fishing effort, exploiting many species at once, induces negative impacts on various species and biological communities in Galapagos (Okey et al. in review). Research elsewhere suggests that a decrease in the populations of exploited species, which in many cases are strong ecological interactors, has cascade or ripple effects on the rest of the marine ecosystem. Data collected in Galapagos suggest that the shallow rocky communities of the Archipelago are significantly altered and unbalanced. A recent assessment of fish communities in the central archipelago showed that artisanal fishing has direct effects on the target species, with flow on effects to community structure (Ruttenberg, 2001). This is the first reported evidence of artisanal fishing producing cascading effects on non-commercial species. That is to say, the ecosystem composition and functioning has changed from its original unexploited state and the food web has also changed, as described in the discussion on Criterion 2 in Chapter 8. Intensive fishing of top predators (groupers, snappers and lobsters) can lead to an increase in populations of grazing invertebrates (particularly sea urchins), which in turn overgraze the substrate. This disturbance impedes the recovery of corals since larvae and recently settled juveniles are unable to establish a foot-hold. In addition, grazers remove most benthic primary production and thus impoverish the biomass of plants and animals on rocky reefs. This is a well-documented ecological phenomenon worldwide (Mann and Breen, 1972; Estes and Palmisano 1974; Glynn et al., 1979; Elner, 1990; Levitan 1992; Sala 1998), and appears to be of widespread occurrence in Galapagos.

Many fisheries affect a range of species, beginning with the removal of top predators. Subsequently the fishing effort is concentrated on species at lower trophic levels resulting in the sequential removal of species from many trophic levels. This practice is referred to as “fishing down marine food webs” (Christensen and Pauly, 1998; Pauly et al., 1999). In recent years there is much evidence that the fisheries effort in Galapagos has diversified to include many species from lower trophic levels, including primary consumers such as sea cucumber (Stichopus fuscus) and certain species of mullet (Mugil galapaguensis and Xenomugil thoburni), but has not yet reached herbivorous fishes or sea urchins.

Growth in fishing sector

Fishing in Galapagos has increased over the years (Figure 9.2). Several pulses of growth in the number of registered artisanal fishers have occurred due to new and more profitable fishing practices or target species. In the early 1940s and 50s, most fishing was done by less than 100 fishers, using hand and drop lines and targeting groupers to produce salt-dried fish for export to the mainland and some limited local consumption. In the early 70s and late 80s, rapid growth was experienced with the development of the lobster fishing for international markets using surface supply diving techniques. In the early 90s another period of growth was experienced mostly due to the sea cucumber fishing (in 1992) and to a lesser extent by illegal shark fin fishing (Bustamante et al. 1999a). Between 1998 and 2000 the number of fishers and fleet size have apparently increased by 92% and 54% respectively, due to the reopening of the sea cucumber fishing for a two-month season, after being closed since 1994. Many of the boat permits issued in 2000 were supposed to be “provisional” and much of the registration of new fishermen with the cooperatives was in contravention of a moratorium limiting new membership to the sons and daughters of fishers. A detailed review in 2001 led to a significant reduction in the number of fishermen but could not withdraw more than a handful of the boat permits. As well as fuelling fleet expansion, the high-value fisheries with Far Eastern markets exacerbates illegal activities and puts great strain on the participatory management process. The prospects of short-term riches can derail long-term programs to achieve sustainable fisheries or to reconcile fisheries with other legitimate interests, such as tourism and biodiversity conservation.

As the numbers of registered fishers increased, the number of targeted species and fishing fleet size rose accordingly, reaching about 1,000 fishers with 444 active fishing boats in 2000 and targeting almost 100 different species of fishes and invertebrates (Fig. 9.2). According to Espinoza et al. (2001), the total wet weight of the catches in 2000 exceeded 667 metric tonnes, and was composed mostly of a variety of fishes (65.3%), sea cucumbers (21.9%) and spiny lobsters (12.8%). Almost 70% of the total catch consisted of 8 species of finfish, lobsters and sea cucumbers, with the most important fishes – mullets (‘lisas’: Mugil galapagensis and Xenomugil thoburni), the serranid grouper (‘bacalao’: Mycteroperca olfax) and scorpaenids (‘brujo’: Scorpaena mystes and Pontinus spp.) – contributing 36% of the total catch (Espinoza et al. 2001).


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Trend on Species and Fishers

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Figure 9.2. Trends of exploited species (open circles), number of registered fishers (grey bars), and numbers of fishing boats (numbers in black) over the past 60 years in Galapagos. Data for the fishers between 1940 and 1960 estimated from personal interviews with older fishers. Number of exploited species in 2000 was considered to be the same as 1999.

In fisheries research situations where population density data independent of the fishery are not available, catch per unit effort or CPUE is widely used as an indicator of species abundance. This indicator needs, however, to be treated with caution, because increasing fishing efficiency or serial site selection may obscure real declines in population numbers. For the Galapagos spiny lobster fishery (Panulirus gracilis and P. penicillatus), CPUE from 1994 to 1997 declined consistently, experiencing thereafter a slight recovery, probably due to strong recruitment after the 1997/98 El Niño (Figure 9.3). Declines in CPUE were suggested to be due primarily to increasing fishing effort and declining adult populations (Bustamante et al. 1999b, Toral et al. 2000). The data from 2001 indicates that CPUE has declined relative to 2000, indicating that the good recruitment year class of 1999-2000 has already largely been removed by fishing and that the downward trend is likely to continue unless total fishing effort can be reduced (Fig. 9.3).

Fishing Trends of Spiny Lobsters

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Figure 9.3. Yearly average catch per unit effort (CPUE) for the 1994-2001 seasons of the spiny lobster fishery of the Galapagos Islands. Boats are artisanal fishing vessels with live-aboard facilities (10-15m length) while dinghies include wooden skiffs or “pangas” and fast open fibreglass boats, both 5-10m length and equipped with outboard engines . Season catches are expressed in metric tonnes (mt). Data for the 19994-96 seasons extracted for summary report of G. Morán (unpublished data) and for 1997-2000 from CDRS Fisheries Monitoring Program.

Although numerous gaps and high variability exist in the data series, catches per season for the hand-line fishery for the Galapagos grouper, Mycteroperca olfax, appears to have been unusually high in 1997/97 and declined to a very low level in 2000/01 (Figure 9.4). As with the lobster population, this could be interpreted as another classical pattern for over-exploited populations. However, the general trend of CPUE remained relatively stable around 0.008-0.022 tonnes/fisher/day (Fig. 9.4), and an increase occurred in 2001 despite the catch having reached an all time low (Fig. 9.4). The drastic declines in catch experienced in the 99-2000 and 2000-2001 seasons could be explained by the fact that little fishing effort has been given to this species and the finfish fishery as a whole since the reopening in 1999 of the sea cucumber fishing in Galapagos (Espinoza et al., 2001). In addition, the whole fin-fishery has changed its behaviour; now given much effort to other species using different techniques (gillnet for mullets, for example) in conjunction with the traditional hand-and-drop line fishing. So, it is likely that the mixing of fishing methods and targets is confusing recent trends in catch and CPUE for these fisheries. Detailed monitoring by on-board observers combined with fisheries independent assessments are needed in order to discern the real trends and status of finfish species.

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Figure 9.4. Total catches in metric tonnes and catch per unit effort for the Galapagos grouper per fishing season.

Another ecologically significant fishery in Galapagos is the use of nets or longline from artisanal or industrial boats to catch sharks for their fins, which are exported to eastern Asia (Zarate, 2002). This lucrative fishery is illegal but has apparently expanded considerably in recent years, due to its profitability. The Galapagos National Park Service is presently attempting to curb this fishery by increasing surveillance.

In conclusion, fishing effort in Galapagos has traditionally been selective for target species and relatively light, but rapid recent growth in numbers of artisanal fishers, particularly new immigrants in search of lucrative sea cucumbers, is putting increasing pressure on coastal food webs. This increasing fishing pressure is likely to interact with El Niño, placing a double strain on the marine ecosystem.


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Pollution

A much smaller and more localised threat to marine biodiversity in Galapagos is domestic pollution and littering in port areas and tourism visitor sites (Bustamante, 1995; Palmer et al. 1999). For the biodiversity vision, this pollution should be considered of minor importance for overall Galapagos marine biodiversity, but an important local issue around human settlements.

A larger threat is the danger of oil pollution from accidental spills, still vividly in the mind from the spill in January 2001. Initial investigations of the effects of the recent Jessica oil spill on marine communities in Galapagos indicated relatively slight impacts, including changes to plant, invertebrate and fish communities within 50 m of the wreck site and sea lions with superficial oiling (Lougheed et al., 2002). Fortuitous ocean currents, which transported oil away from and between islands, and high levels of solar radiation, which accelerated evaporation of volatile petroleum compounds, contributed to the low apparent levels of impact. Nevertheless, a long-term study of marine iguanas by Wikelski and colleagues, which includes good historical information for the affected island of Santa Fe, indicates substantive population effects of oiling. The marine iguana population under study showed signs of stress immediately after the oiling, at a level normally associated with high mortality (Wikelski et al. 2001). Data from November 2001 indicates apparently high mortality in this study population, probably as a consequence of the contamination (Wikelski et al. 2002). Flightless birds, such as the Galapagos penguin and flightless cormorant, are especially vulnerable to oiling, but in this case appear to have been unaffected, as they were far from the spill. The studies following the Jessica oil spill highlight the importance of creating a good baseline data set, in order to assess human impact and monitor change over the medium- to long-term.

Tourism

Apart from pollution effects, tourism in Galapagos, being a non-extractive activity, has little impact on the marine ecosystem, in part because the environment is mostly rocky reef rather than coral. Studies at a heavily used snorkelling site detected no significant impacts (Monsalve and Bustamante, 1997, ). Of course, if tour boats catch fish, as many have been doing over the years, then this activity has the same impact as any other fishing activity, commercial or not, at the same scale in the same location – the impact on local fish is the same regardless of who is doing the fishing. However, regulations are now being put into effect to curb fishing by tourism boats (Arboleda and Montoya, 2000). On the other hand, catch-and-release sport fishing is presently being discussed and may commence soon in Galapagos.

Introduced species

In comparison to the impacts of fishing, pollution and tourism, which can be ameliorated through adaptive management, the threat of introduced species is a long-term one that will continually increase rather than decrease over time. Introduced species thus represent much more of a potential than a realised threat to local Galapagos marine biodiversity.

No water-ballasted cargo boats presently operate to Galapagos, hence introductions through ballast water are unlikely; however, marine species can be dispersed to Galapagos through hull fouling and deliberate human translocation. Species arriving in Galapagos as fouling organisms are poorly known because of their cryptic nature, but possible candidates include species of barnacles, hydroids, bryozoans, serpulid worms, sponges, bivalves, and ascidians. Species deliberately or accidentally introduced by direct human transport have occurred in the case of the mangrove crab Cardisoma crassum, a crab brought to the islands deliberately for human consumption in the early 90’s and now known to exist in the mangroves on Santa Cruz (C. Marquez pers. comm.). The ‘chame’, Dormitator latifrons, an estuarine fish abundant in the mainland, now dominates coastal lagoons on the island of Isabela. It is not know how this species got to the islands, but it could well turn out that this is a native species, since very little attention has been given to the biodiversity of freshwater and coastal lagoon of the Galapagos.

All boats, including tour boats, can contribute to the dispersal of certain species around the archipelago, and to the risk of introducing alien species through waste disposal or hull fouling. Because of their wide-ranging itineraries, ocean-going yachts and cruise ships represent a particular risk in terms of the introduction of invasive alien species, terrestrial as well as marine.


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9.4 Status of Marine Biodiversity According to the Three Criteria

The following three tables summarise the current status of marine biodiversity, as measured by the three criteria defined in Chapter 8, relative to the supposed pristine state. For many of the indicators, especially those under Criterion 2 that deal with ecosystem composition and functioning, few data for Galapagos are available. However, the underlying processes are known for other marine ecosystems and it is assumed that Galapagos ecosystems are not very different.

CRITERION 1 – Extent to which biogeographic zones and habitats are represented in No-Take-Areas (NTAs) and surrounded by areas with effective management. Indicator 1999 status relative to

“pristine” state Conservation opportunities and threats

Proportion of each habitat in each biogeographic zone in NTAs

0%. The 1992 management plan for the former Marine Resources Reserve defined and mapped NTAs but these were never applied in practice.

Proportion of coastal areas within effectively protected NTAs

0%

Opportunity: The 1998 Special Law for Galapagos (SLG) creates a new Marine Reserve, including many habitat types, and requires that it be zoned. The 1999 Management Plan defines categories of No-Take-Areas (NTAs) and a process for establishing first provisional then permanent zonation over a period of 4 years. The provisional zonation was approved in April 2000 and came into effect immediately, but final definition and delimitation of zone boundaries was delayed; it is scheduled for early 2002. NTA’s will occupy just over 17% of the coastline but include no open ocean areas. Threat: NTAs may be too small and too few, and may not be respected by either fishermen or the local tourism boats, which have been accustomed to fish. The control and enforcement capacity of authorities may not increase sufficiently or cover the total NTAs network.

Proportion of NTAs surrounded by well managed areas

0% Opportunity: The Special Law for Galapagos (SLG) and complementary norms for fishing and tourism, if well enforced, makes it possible to control human uses and could effectively put a cap on fishing expansion and control impacts. Threat: Appropriate control may not occur, resulting in over-exploitation of several Galapagos fisheries. Fisheries management capability in the islands may be slow to build up, as may enforcement capability. Other impacts, such as pollution, are localised and minor, but if human immigration continues and sources of pollution are not controlled, including the frequency of supplies by boats and planes, pollution could pose a serious threat to marine biodiversity, including the arrival of marine aliens (biological “pollution”). Oil spills are an ongoing threat.


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CRITERION 2 – Extent to which composition and functioning of the ecosystem are maintained. Indicator 1999 status relative to


Status of selected taxa indicating ecosystem health (priority species only)

Sea urchins and herbivorous fish are abundant. Seaweeds, black corals, marine iguanas, predatory fish (groupers, snappers, jacks), lobsters, sea cucumbers, sharks, predatory snails and some cetaceans are all significantly reduced by human activities. Surveys in 2001 suggest pinnipeds populations also reduced, perhaps by El Niño combined with human activities. Barnacles disappeared in 1997/98 El Niño but are recovering as part of natural cycles. Seaweeds, hermatypic corals and gorgonians also affected by El Niño. Some seaweeds and associated communities appear to have been lost since 1983 El Niño. Overall ecosystem functioning and composition, including its trophic structure, is altered.

Species diversity and abundance

Species diversity and abundance are moderately disturbed. Few known extinctions, but few data on rare species.

Opportunities: NTAs network (zoning) could quickly protect substantial parts of habitat and exploited species allowing recovery of most taxa. Quick responses are expected after NTAs implementation (<5-10 years). NTAs spillover effects are predicted and NTAs will hopefully gain support from fishers and tourist operators. Industrial fishing has been prohibited. Threats: Exploitation, which has increased in recent years, and its interaction with El Niño, are the main threats. For coastal marine species, introduced species are a threat, e.g. cats, rats, dogs and pigs prey on marine iguanas, nesting sea and shore birds and sea turtles. Sperm whales do not seem to be recovering, despite local protection. Seaweed distribution, biomass and species richness have all declined, possibly because of depletion of fish that prey on grazers. Fisheries for lobster, sea cucumber and predatory fish appear to be unsustainable at current levels and the trend is to broaden the range of exploited species. There is pressure to permit shark as “by-catch” of the developing offshore tuna and billfish fishery, and also to start a sea urchin fishery. Locally based pelagic fisheries, which are to be encouraged as an alternative to coastal fisheries, may have by-catch problems affecting turtles, pinnipeds, seabirds etc. Global climate warming means that the ENSO cycles will be occurring on top of an increased baseline temperature, so that the El Niño periods may be increasingly intense. As well as having a direct impact on many marine species that have low tolerance to El Niño conditions, there is also a risk of synergy between these effects and those of increased fisheries, causing further destabilisation of the ecosystem.

Genetic diversity in populations at risk

Some populations are naturally small e.g. penguins. Others have been through bottleneck e.g. fur seals, sperm whales. Some are subject to directional selection – lobsters, bacalao (grouper). But there are no genetic data.

Opportunity: Legal framework for enhanced protection to endangered and species at risk, especially to those endemics of low abundance. Threat: Several land-based marine species continue in low numbers. Penguin population remains low for unknown reasons. Selection on commercial species is intensifying.


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Isolation and alien species

Still relatively isolated. Average endemism around 20%. Marine alien species not yet a significant problem, as far as is known.

Opportunity: There are no marine alien pest species recorded to date (though one species of crab has been reported in Santa Cruz). The SLG created the Galapagos quarantine inspection system that should be used, with Navy support, to extend controls to marine risks. Most cargo boats come to islands loaded and return empty, minimizing the risk of ballast water introducing aliens. Threat: Increasing boat traffic to and within islands (passenger and cargo transport, tour boats, fishing boats, ocean-going yachts) poses a threat, through waste disposal and alien species “hitch-hiking” attached to boat hulls or in ballast waters. As notoriously invasive marine species spread around the world, the risk of them reaching Galapagos increases.

Pathogen and parasite prevalence

Little information, but suspected avian pox of seabirds and possible natural outbreaks of sea lion parasites (blinding pups).

Opportunity: To date there is no record of (or research on) marine diseases nor has there been any aquaculture. Threat: Isolation has made Galapagos species potentially vulnerable to pathogens. Sea lions can pick up dog pathogens (there was an outbreak of canine distemper in Galapagos dogs in April 2001). Penguins may be at risk. Proposals to develop aquaculture pose a disease threat, either by introduction of pathogens or by development of high -density cultured populations.

Resilience of biological communities

Response after El Niño seems variable, but data are limited. Penguins, barnacles, at least two starfish, seaweeds and corals have failed to recover fully from 1983 El Niño. Sea cucumbers have not recovered from exploitation, but this has never really stopped.

Opportunity: Recent analyses have overwhelmingly demonstrated the effectiveness of marine reserves in protecting and restoring marine biological communities altered by fishing. The new zoning scheme of the GMR could represent a viable means of monitoring natural variation and climate change while allowing recovery of exploited species. Threat: Greatest threat may be the interaction between the extreme natural variability of the Galapagos marine environment and anthropogenic impacts, principally exploitation and alien species. If El Niño events intensify, then the risk of irreversible change due to such interactions will increase.

Connectivity/ recruitment

Few data on recruitment or larval dispersal to guide management. Lobster and sea cucumber stock-recruitment relation poorly known. Ongoing genetic work may shed some light.

Opportunity: The isolation, varied sizes of islands, the relatively small distances and the biogeographic heterogeneity of Galapagos marine ecosystem make it on of the best places to test recruitment dynamics (sink-source and retention-dispersal patterns). The layout of the NTAs network (sizes, numbers, distances), represents one of the best marine sites to verify the roles of the indicators of connectivity and recruitment. Threat: Since we know little about connectivity in Galapagos (or anywhere else in the world), we cannot identify specific threats. Meanwhile precautionary approach needed.


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Habitat modification

Localised habitat modification and pollution close to the 5 ports. Some beaches badly affected by removal of sand for building. No known region-wide modification. Oil spill from tanker, Jessica, caused widespread pollution in 2001.

Opportunities: Jessica incident served as a wake-up call for better regulation of shipping, contingency planning and investment in pollution control capability. Donor support is likely for systems to reduce sewage pollution in port areas. Threat: Increasing human population could increase local impacts. Regional modification only likely as result of exploitation or alien species. There is a risk of oil spills; no contingency plans exist.

Land-sea interactions

Few data, but natural interactions involving seabirds, mammals, and reptiles largely maintained. Terrestrial alien species are the one major problem. No major rivers and only localised pollution sources.

Opportunity: Controls on terrestrial development make it very unlikely that land-sea interactions will be disrupted, other than by alien terrestrial species (discussed above). Programs for control of alien species are being strengthened. Threats: Impacts of alien species on marine species, whilst they are on land, could increase. Fisheries could affect seabirds, mammals and reptiles through by-catch and competition. Iguanas impacted by dogs and cats.

CRITERION 3 – The value and sustainability of products and services. Indicator 1999 status relative to


Abundance and population structure of large marine animals

Some species have declined (e.g. whales and sharks). Sperm whales do not seem to be recovering, despite protection. Sharks are under intense pressure from illegal fishing by local and external fishermen. Fur seals have recovered from near extinction in late 19th Century but recent estimates suggest total population may be small: perhaps as low as 6,000 individuals (Salazar, 2002).

Opportunity: Large marine fauna still found in numbers that allow use by tourism and science. Sharks have a high touristic value as the flagship species of Galapagos dive tourism. If no protection is given elsewhere, the marine reserve may represent the only refugia for wide-ranging and resident populations. The monitoring of population structure and abundance for the large marine fauna is a costly activity, but the continuous presence of tour boats provides an opportunity for participatory data collection and ecological monitoring. Threat: Continuing shark-fin fishing, industrial fishing and pollutants from remote sources around the reserve and in the region may well still cause negative impacts for the wide ranging megafaunal species (sharks, whales, sea turtles) through bio-accumulation. Some fishing methods, such as longlining, could affect pinnipeds, turtles, sharks etc. Pinnipeds have also reportedly been killed as shark bait because fishermen view them as competitors, or because there is a market for their penises as aphrodisiacs.


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Abundance and population structure of seabirds

Most probably remain near original levels. Dark-rumped petrel rescued from trend towards extinction but still far from original abundance. Penguins have not recovered fully from decline during El Niño of 1983.

Opportunity: Local coastal fisheries have little impact on seabird species. Also, the limited development of offshore fishing could be a positive factor for the development of alternative and low impact fishing technologies that will result in the reduction or mitigation of the by-catch. Significant advances and good prospect of controlling introduced predators. Threat: There is little management or control of the impact of the coastal and offshore fisheries using small-scale intensive fishing gears prone to by-catch (long-lines, gill nets). If El Niño’s become more intense, many seabird species will suffer and rare ones may face extinction (especially penguin).

Abundance, size and population structure of reef fish

From original big schools and large size, large predator species (especially groupers) have declined severely in abundance and size. Some lower trophic level fish also reduced.

Opportunity: Recovery of few large predators could be achieved with the NTAs network since there are still some remaining stocks and juveniles are present in most reefs. Spearfishing is banned and gillnetting severely restricted. Threat: All reef fish species lack management or regulation. Some species may continue to be exploited and will not recover, due to slow growth, erratic or minimal recruitment, or the inadequacy of the network of NTAs.

Number and mean trophic level of exploited species

Originally groupers were main target species and < 30 species were exploited. Now groupers less common in catch and species of lower trophic level exploited. Total number of species caught > 90. Marked declines in populations of lobster, sea cucumber, groupers, sharks.

Opportunity: Artisanal fisheries have operated so far on few target species of top predators, while most of the populations of mid-to-small size lower trophic predatory and herbivorous reef fishes have not sustained commercial exploitation. Threat: The once-abundant high trophic level species (groupers, snappers and lobsters) have not shown any sign of recovery and fishing is still at high levels (as are market demand and price)

Catch Per Unit Effort for exploited species

In the past 1-2 decades CPUE has declined for lobster, sea cucumber and some fish species.

Opportunity: Low availability of the target species and consequently low economic returns, if prices and demand do not increase, could lead to a reduction of overall fishing effort if economic alternatives are available. The effective implementation of a network of NTAs could enhance fishing grounds and maintain CPUE values. Threat: If reduction of fishing effort is not achieved, stock rebuilding policies are not in place, and there is no rapid effect of NTAs network on improving catches, then CPUE is expected to decline further.

Spawning stock of exploited species

Probably low in some species e.g. sea cucumber.

Opportunity: The effective implementation of a network of NTAs should protect a fraction of the spawning stock, allowing the possible recovery of exploited species. Threat: The exploited species within the NTAs network may not recover quickly, due to any combination of the following factors: NTAs (directly related to protected spawning stock) may not be large enough; the response is not fast enough (<2-4 years); the impact of exploitation outside of the NTAs is still growing and unmanaged; poaching in NTAs.


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Indirect impacts, such as by-catch, pollution, physical damage to corals

Originally there were no such impacts. Now increasing by-catch of seabirds, mammals, sharks and other large animals reported for offshore fisheries but not measured. Localised waste and nutrient pollution. Some tour operators already adopting pollution control measures. Little anthropogenic damage to corals.

Opportunity: The indirect impacts are probably of low magnitude, so mitigation, reduction or elimination is still possible with sound management. Threat: No management provisions for by-catch have been developed and, even if they were, there may be problems of compliance. Some fishermen aspire to semi-industrial fishing methods likely to cause by-catch problems and continue to press to be allowed to use “artisanal” longline in the GMR. Other indirect impacts may increase as human activities (fishing, tourism, development) increase.

Benefit relative to volume of extraction

Marine tourism and fisheries both on increase. Fisheries catch has broadened to include more species, mostly of lower value, and to smaller individuals. Export market for spiny lobster has kept prices up. Sea cucumber price paid to fishermen was high but dropped in 2001. Trend in total benefit relative to volume of extraction unknown.

Opportunity: Galapagos potential for marine tourism so far little exploited. Galapagos has a strongly conservation-oriented tourist industry that is an example to the world. Financing for change from extractive to non-extractive activities could be available, if a permanent reduction in fishing fleet were consequently assured. Threat: Sea cucumber, shark fin and lobster are high value products, so sustainable, non-extractive alternatives cannot compete with intensive, unsustainable fisheries. Furthermore, few fishers may want to convert to non-extractive activities, because of lack of skills and loss of independence.

Level and orientation of subsidies

Fees for tourism in Park subsidise fisheries management; fisheries sector pays no user fees. Remaining management and research costs covered by donors and government. Fuel subsidy, which stimulates both fishing and tourism, is being reduced. Prevalence of tax evasion, which has similar effects to a subsidy, in Galapagos tourism and fisheries sectors is unknown.

Opportunity: There is a general national trend to reduce subsidies and to tighten up on taxes. Local fishermen accept that the fisheries sector, including traders, should contribute to management costs of sea cucumber. In 2001 an attempt was made to have traders contribute 4 cents per sea cucumber to the fishing cooperatives and to management costs; most paid up but the cooperatives kept all the money. Perverse subsidies encouraging unsustainable fishing should cease. Threat: There is not yet a legal mechanism for charging fees for fishing.

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CDF/WWF Biodiversity Vision Chapter 10 – Marine Biodiversity Vision

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CHAPTER 10 – PROJECTIONS FOR THE FUTURE: A MARINE BIODIVERSITY VISION

Principal Authors – R. Bensted-Smith, G.M. Branch, R.H. Bustamante, and G.M. Wellington.

Assumptions

The projections presented in this chapter for marine biodiversity in the year 2050 are based on three sets of assumptions: business-as-usual, improved but not ideal management and optimal management. The latter projection generates the biodiversity vision.

The business-as-usual scenario assumes that the present management regime continues its efforts but the major obstacles to conservation continue to impede conservation efforts. Thus the principal assumptions would be:

• Continuing expansion of the number of fishers and capacity of the local fishing fleet;

• Regulations do not provide an effective mechanism to restrict or even reduce fishing capacity;

• Poor compliance with the laws and limited effectiveness of patrolling and prosecution;

• Management plan acknowledged and NTAs recognised by fishers but illegal fishing is frequent in most locations;

• Participatory management system functioning but dogged by controversies over actual or potential high value fisheries; internal organization of participating sectors improves slowly;

• Community awareness and activism in favour of conservation increases slowly;

• Scientific information continues to be used in decision-making but is often over-ridden where there is pressure from fishers;

• Marine management capability and financial resources of GNPS and the effectiveness of its collaboration with the Navy improve slowly;

• Research and monitoring capability and financial resources of CDRS experience fluctuations, according to fund-raising success;

• Continuing external threat from industrial fishing fleet;

• International attention from UNESCO and other organizations continues.

• El Niños occur periodically, with a moderate increase in intensity due to global climate change.

The derivation from this list of the assumptions for the optimal management scenario is straightforward. However, the group also decided to make a projection based not on business-as-usual but on an assumption that several of the major obstacles will be tackled in the coming year or two. The justification for this is that the legal, institutional and managerial framework for the GMR is in a state of rapid development, as is the social and political environment for marine conservation. Furthermore, there are prospects for Inter-American Development Bank investment. Therefore, this “improved management” scenario is based on the following intermediate assumptions, the validity of which up to early 2001 is indicated in parentheses:

• The Special Regulation for Fisheries in Galapagos is satisfactory and is promulgated in 2000. (It was near finalisation in late 2001 but in 2002 approval has been delayed and the draft regulation weakened. It may nevertheless include a number of positive measures, including a limit on the size of the local fishing fleet. Unfortunately, in 2000 the Inter-


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institutional Management Authority, under aggressive pressure from the fishing sector, authorised the issuing of some 190 new fishing permits for local boats on a provisional basis. These were reviewed in 2001 but almost all have been ratified. This expansion will make it more difficult to achieve sustainability and will increase conflicts between fisheries and conservation, because the benefits of sustainable fisheries must be shared between more people. Indeed, such conflicts erupted in the lobster fisheries in November 1999. On the other hand, all sides, including the fisheries sector, now recognize the need to reduce the local fishing fleet.)

• The participatory management system is substantially strengthened. (Work continues satisfactorily despite occasional setbacks, such as the fisheries conflicts in 2000.)

• Active community support for marine conservation increases markedly. (Work continues; some supporting groups are becoming increasingly active but there are other groups that generally oppose conservation measures.)

• GNPS marine management capability is substantially strengthened during the next five years by external investment. (Limited external support will soon be complemented by a substantial Inter-American Development Bank loan. GNPS has invested own resources in marine management strengthening.)

Further comments about the assumptions are contained within the tables below, which describe the projections in terms of the three Criteria identified earlier. The information on 1999 status is repeated for ease of reference.


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CRITERION 1 – Extent to which biogeographic zones and habitats are represented in No-Take-Areas (NTAs) and surrounded by areas with effective management. INDICATOR STATUS 99 BUSINESS AS USUAL IMPROVED MANAGEMENT (1) VISION Proportion of each habitat in each biogeographic zone in NTAs

0% The 1992 management plan for the former Marine Resources Reserve defined and mapped NTAs but these were never applied in practice.

Habitats (in each biogeographic zone): 10% of coastal habitats in NTAs, with some habitat types under-represented e.g. mangroves and sandy beaches. Some biogeographic zones also under-represented e.g. northern. 0% of offshore habitats, including sea mounts, in NTAs.

0-50%: some habitats wholly outside NTAs. Rocky, exposed – 15% Sandy beaches – 30% Lagoons – 60% Mangroves – 60% Salt marsh –10 % Coral reefs –50 % Shallow subtidal reefs, exposed – 15% Shallow subtidal reefs, sheltered – 15% Sea mounts – 5% Offshore slopes, rocky – 5% Offshore slopes, sand – 5% Plains > 1000m – 5% Pelagic nearshore – 5% Pelagic offshore – 5%

16 major habitats protected 20-100% by NTAs in each biogeographic zone where they appear. Rocky, exposed – 20% Sandy beaches – 100% Lagoons – 100% Mangroves – 36% Salt marsh – 36% Coral reefs – 50% Shallow subtidal reefs, exposed – 36% Shallow subtidal reefs, sheltered – 36% Sea mounts – 36% Offshore slopes, rocky – 20% Offshore slopes, sand – 20% Plains > 1000m – 20% Pelagic nearshore – 20% Pelagic offshore – 20%

Proportion of coastal areas within effectively protected NTAs

0% 10% of coastline within NTAs but with various exceptions to the rules and poor protection, so that only the core areas of large NTAs are really free of extraction.

20% of coastline within NTAs, with fairly effective protection, except in some small, remote NTAs. Some initial exceptions, such as allowing fishing for mullet at certain beaches, are gradually closed down.

About 40% of the coastline in NTAs, all fully protected. The proportion would be higher for scarce, sensitive habitats e.g. coastal lagoons, coral reefs and sandy beaches.

(cont)


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INDICATOR STATUS 99 BUSINESS AS USUAL IMPROVED MANAGEMENT (1) VISION Proportion of NTAs surrounded by well managed areas

0%

0% Fishing continues to intensify and tends towards the semi-industrial. Some of it is controlled but nevertheless unsustainable. New species are exploited. Illegal industrial fishing and also illegal local and external fishing of vulnerable, high-value species, such as sharks, continue. Some species, such as sea cucumber, become commercially extinct. There is intense pressure to open NTAs to fishing.

50% Illegal fishing is reduced. Legal fishing extends to one or two more species and there is over-exploitation of some benthic species. However, the expansion of fisheries levels off, through regulations to limit fishing effort and alternative economic and educational opportunities. Outside the GMR the problem of unmanaged industrial fishing continues.

95% There is good management throughout the GMR, with no unsustainable exploitation and negligible illegal fishing, so only the species that move outside the GMR boundaries would be inadequately protected. Fishing effort is well distributed between species and is strictly artisanal. Fishing effort is gradually reduced, due to firm regulation of permits and status as “fisherman”, combined with imaginative training and incentive schemes for conversion to non-extractive activities.

(1) Crude % estimates. New calculations are expected using a complete set of satellite images and GIS analyses.


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CRITERION 2 – Extent to which composition and functioning of the ecosystem are maintained. INDICATOR STATUS 99 BUSINESS AS USUAL IMPROVED MANAGEMENT VISION Status of selected taxa indicating ecosystem health (priority species only) For details of the taxa and their status, vision etc see Annex 8.1

Sea urchins and herbivorous fish are abundant. Seaweeds, black corals, marine iguanas, predatory fish (groupers, snappers, jacks), lobsters, sea cucumbers, sharks, predatory snails and some cetaceans are all significantly reduced by human activities. Surveys in 2001 suggest pinnipeds populations also reduced, perhaps by El Niño combined with human activities. Barnacles disappeared in 1997/98 El Niño but are recovering as part of natural cycles. Seaweeds, hermatypic corals and gorgonians also affected by El Niño. Some seaweeds and associated communities appear to have been lost since 1983 El Niño. Overall ecosystem functioning and composition, including its trophic structure, is altered.

Sea urchins and herbivorous fish decline, due to future exploitation. Pinnipeds continue natural fluctuations; but may decline, as may cetaceans. Marine iguanas decline further, due to El Niño fluctuations interacting with anthropogenic factors. Predatory fish, lobsters, sea cucumbers, sharks and predatory snails all decline further due to exploitation. Seaweeds, gorgonians and black corals maintain present levels. Hermatypic corals recover somewhat, but slowly. Barnacles and ascidians continue to fluctuate naturally. Trophic structure becomes increasingly modified.

Sea urchins and herbivorous fish populations maintain present levels. Pinnipeds continue natural fluctuations but may decline, as may cetaceans and sharks, unless oceanic no-take areas are established as well as the coastal NTAs currently being established. Marine iguanas will continue to fluctuate with El Niño but may not decline. Populations of predatory fish, lobsters and predatory snails may recover locally in numbers and size. Sea cucumbers may or may not recover– we know too little about their population biology. Seaweeds and gorgonians maintain present levels, hermatypic and black corals recover slowly. Barnacles and ascidians continue to fluctuate naturally. Trophic structure becomes more like natural state.

Data will be available on all these indicator taxa. For indicator species in general, populations maintain natural levels, demography and cycles. For seaweeds and hermatypic corals areas of high diversity of these taxa are fully protected. Localised nutrient pollution problems are controlled. Certain taxa are completely free from exploitation: herbivorous fish (a key guild in ecosystem), cetaceans, pinnipeds, sharks, predatory snails, black coral. Populations of some taxa are restored to pre-exploitation levels: sea cucumber, predatory fish, lobsters, predatory snails, black coral. Cetaceans will take more time but should recover in an enhanced pelagic environment. Marine iguanas populations are also restored (requires control of terrestrial predators). Trophic structure reverts fully to natural state.

Species diversity and abundance

Species diversity and abundance are moderately disturbed. Few known extinctions, but few data on rare species.

Increasing levels of disturbance of patterns of diversity and abundance, with populations of some species reduced to minimal levels. Commercial extinction of some species.

Disturbance increases but less than if business as usual. More data are available on the abundance and distribution of species. No human-induced extinctions.

Minimal anthropogenic impact. Comprehensive data are available on diversity, abundance and distribution of species.


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INDICATOR STATUS 99 BUSINESS AS USUAL IMPROVED MANAGEMENT (1) VISION Genetic diversity in populations at risk

Some populations are naturally small e.g. penguins. Others have been through bottleneck e.g. fur seals, sperm whales. Some are subject to directional selection – lobsters, bacalao. But there are no genetic data.

There is no genetic data but a loss of diversity in exploited species is presumed.

Genetic data available on a few species. There is a loss of diversity in exploited species, but effect moderated by NTAs.

Genetic information available on a range of species. Natural levels of genetic diversity maintained or restored (a slow process that would be barely starting in 50 years).

Isolation and alien species

Still relatively isolated. Average endemism around 20%. Marine alien species not yet a significant problem, as far as is known.

Ecosystem further modified, including presence of alien species. Mariculture introduced.

Some introduction of alien species, principally through boat traffic. Mariculture a possibility.

All endemic species preserved. Minimal modification by alien species. Mitigation or restoration, where possible. Mariculture not introduced.

Pathogen and parasite prevalence

Little information, but suspected avian pox of seabirds and possible natural outbreaks of sea lion parasites (blinding pups).

Unknown. Unknown. Anthropogenic introduction of pathogens and parasites prevented by regulations on shipping, avoiding introduction of mariculture, and other measures. Data available on pathogens and parasites of selected species.

Resilience of biological communities

Response after El Niño seems variable, but data are limited. Penguins, barnacles, at least two starfish, seaweeds and corals have failed to recover fully from 1983 El Niño. Sea cucumbers have not recovered from exploitation, but this has never really stopped.

There is the risk that biological communities will be altered so much that they lose resilience. Spawning stocks may be reduced to levels at which fertilisation fails (Allee effects).

Resilience is likely to be maintained, because of connectivity between network of NTAs and because of reduced disturbance in general. However, it is not known if some biological communities would withstand El Niño phenomena of increased intensity due to climate change.

Resilience is maintained. Reduction of synergistic interactions between disturbances. Management interventions are taken to respond to anthropogenic disturbance, thereby assisting natural resilience.

(cont)


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INDICATOR STATUS 99 BUSINESS AS USUAL IMPROVED MANAGEMENT (1) VISION Connectivity/ recruitment

Few data on recruitment or larval dispersal to guide management. Lobster and sea cucumber stock-recruitment relation poorly known. Ongoing genetic work may shed some light.

Unknown. Unknown. Understand connectivity and recruitment patterns of priority species. Establish effective NTAs that maintain (or restore) natural connectivity between areas and natural recruitment patterns.

Habitat modification

Localised habitat modification and pollution close to the 5 ports. Some beaches badly affected by removal of sand for building. No known region-wide modification. Oil spill from tanker, Jessica, caused widespread pollution in 2001.

Increased habitat modification and pollution (land-based and from boats). Oil spills increase.

Trend to habitat modification is stopped, with restoration where NTAs established. Boat pollution reduced. Increased availability of information on habitat types, characteristics and distribution.

Restoration or partial restoration of habitat in all affected areas, due to changes in human activities through education and regulation. Pollution minimised; no lasting impacts. Comprehensive information available on habitat types, characteristics and distribution. Oil-spill contingency plans developed and infrastructure available for implementation.

Land-sea interactions

Few data, but natural interactions involving seabirds, mammals, and reptiles largely maintained. Terrestrial alien species are the one major problem. No major rivers and only localised pollution sources.

Natural interactions are increasingly disrupted due to changes in terrestrial or marine environment or direct interference by human activities (land and sea traffic, infrastructure, extraction, lighting, noise, disturbance etc).

As a result of improved management of human activities, disruptions of natural interactions do not increase significantly.

Natural interactions understood and maintained.


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CRITERION 3 – The value and sustainability of products and services. INDICATOR STATUS 99 BUSINESS AS USUAL IMPROVED MANAGEMENT VISION Abundance and population structure of large marine animals

Some species have declined (e.g. whales and sharks). Sperm whales do not seem to be recovering, despite protection. Sharks are under intense pressure from illegal fishing by local and external fishermen. Fur seals have recovered from near extinction in late 19th Century but recent estimates suggest total population may be small: perhaps as low as 6,000 individuals (see Chapter 9).

Marine Reserve helps to regulate fisheries but pressures on high-value species increases shark harvesting. Sea mammals suffer from increasing by-catch. Community structure is permanently altered. Whale populations fail to recover.

Improved regulations and control protect sharks from deliberate industrial and artisanal fishing. However, they and other large animals continue to suffer from by-catch losses due to increased artisanal fishing.

Stocks are built up again to a level corresponding to zero exploitation. In the case of the whales, this would take much longer than the time-scale considered here. By-catch of large animals zero or negligible.

Abundance and population structure of seabirds

Most probably remain near original levels. Dark-rumped petrel rescued from trend towards extinction but still far from original abundance. Penguins have not recovered fully from decline during El Niño of 1983.

Penguins may go extinct, through a combination of climate change and local anthropogenic factors (introduced predators, disease, change in prey abundance due to fishing). Dark-rumped petrel populations still at risk due to low survival at sea. Species such as waved albatross, boobies and swallow-tail gull reduced in numbers due to increasing by-catch losses.

No extinctions, unless increasingly intense and frequent El Niño’s combine with anthropogenic factors to push penguins into terminal decline. By-catch losses are reduced but continue to represent a significant problem for seabirds, which depend on high adult survivorship.

Seabird abundance restored. Breeding sites sustainably protected. No industrial or semi-industrial fishing or other impacts on small fish. No by-catch of seabirds. Domestic predators (dogs, cats) eliminated or controlled; spread to unoccupied islands prevented.

(cont)


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INDICATOR STATUS 99 BUSINESS AS USUAL IMPROVED MANAGEMENT (1) VISION Abundance, size and population structure of reef fish.

From original big schools and large size, large predator species (especially groupers) have declined markedly in abundance and size. Some lower trophic level fish also affected.

Groupers etc continue their decline. Species of lower trophic level follow a similar trend as fishing pressure continues to grow. Few large individuals remain.

Expansion of fisheries fleet levels off and spear-fishing curbed. Grouper size and abundance remains low, but populations of lower trophic level species do not suffer a similar decline. NTAs provide sanctuaries where populations are restored.

Age and size structure of all reef fish populations restored. Fishing pressure is reduced and is strictly legal, artisanal fishing. Good management of ecosystem and of key species aids restoration. NTAs effective and spill-over of adults and larvae enhances stocks fished outside NTAs.

Number and mean trophic level of exploited species

Originally groupers were main target species and < 30 species were exploited. Now groupers less common in catch and species of lower trophic level exploited. Total number of species caught > 90. Marked declines in populations of lobster, sea cucumber, groupers, sharks.

Number of fish species exploited increases and mean trophic level of catch declines.

Trend towards lower trophic levels is arrested, but high fishing pressure prevents restoration to a better trophic balance. Historic trophic balance restored in NTAs.

Stable fishery, balanced trophically and based on fewer species than at present, sustains local human population, including tourists.

Catch Per Unit Effort for exploited species

In the past 1-2 decades CPUE has declined for lobster, sea cucumber, and some fish species.

CPUE continues to decline as stocks decrease and the local fleet expands. New species entering the fisheries follow the same trend.

CPUE is stabilized and even increases slightly for lobster and sea cucumber, due to management measures. CPUE for reef fish is stabilized as fleet capacity is stabilized and offshore fisheries are developed.

CPUE increased for all species, as fleet capacity and effort decline and as good management of the ecosystem and of target species increases stocks.

Spawning stock of exploited species

Probably low in some species e.g. sea cucumber.

Decline of spawning stock of high value species (sea cucumber, lobster, shark) continues, in some cases to commercial extinction. Other species (e.g. sea urchin) are exploited and follow the same trend, due to intense demand from local fishermen.

For lobster and sea cucumber, control of fisheries is strengthened and management of the species improves steadily, due to better data and growing commitment of fishermen to sustainability. Shark fishing, which is illegal, ceases.

Spawning stock is at least 50% of unexploited level for all species. Sites important for spawning are protected.

(cont)


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INDICATOR STATUS 99 BUSINESS AS USUAL IMPROVED MANAGEMENT (1) VISION Indirect impacts, such as by-catch, pollution, physical damage to corals

Originally there were no such impacts. Now increasing by-catch of seabirds, mammals, sharks and other large animals reported for offshore fisheries but not measured. Localised waste and nutrient pollution. Some tour operators already adopting pollution control measures. Little anthropogenic damage to corals.

By-catch becomes an increasingly serious problem, as illegal industrial fishing continues and artisanal offshore fishing increases and tends towards semi-industrial. Pollution from towns stabilised or reduced, due to investment in treatment. Pollution from tour boats unchanged but other boat traffic (cargo, local transport) increases. Sewage-related diseases start to appear. Visitor impacts may affect rate of recovery of coral structures.

Industrial fishing curbed but by-catch continues to be a problem for some species, as artisanal offshore fishing increases. Pollution from towns reduced. Pollution from tour boats reduced as move to “green” operations grows. Contingency plans prepared for serious spills.

Minimal by-catch within the Marine Reserve. Garbage and other waste properly managed. Reef damage by anchors or underwater tourism prevented. Contingency plans for oil spills developed and backed by adequate funding and infrastructure.

Benefit relative to volume of extraction

Marine tourism and fisheries both on increase. Fisheries catch has broadened to include more species, mostly of lower value, and to smaller individuals. Export market for spiny lobster has kept prices up. Sea cucumber price paid to fishermen was high but dropped in 2001. Trend in total benefit relative to volume of extraction unknown.

Fishing effort continues to grow as fishermen invest profits in boats and equipment. Trends towards lower value species and smaller individuals continue. Marine tourism growth impeded by declining populations of sharks and other attractive big animals.

Fishing effort and exploitation patterns stabilized. Dive tourism grows, as Galapagos maintains populations of big attractive animals, which are ever scarcer in the world. Some fishermen opt for training to shift to non-extractive activities, but it is not easy for them, nor does it significantly reduce the fishing fleet capacity.

Tourism and fisheries mutually sustained, providing increased, widely distributed benefits. Extractive uses do not detract from the non-extractive. In particular, the reputation of Galapagos as the best place to see large marine animals is fully established, providing it with a very robust, high value market. Local use and price of fish rises due to high tourist demand.

(cont)


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INDICATOR STATUS 99 BUSINESS AS USUAL IMPROVED MANAGEMENT (1) VISION Level and orientation of subsidies

Fees for tourism in Park subsidise fisheries management; fisheries sector pays no user fees. Remaining management and research costs covered by donors and government. Fuel subsidy, which stimulates both fishing and tourism, is being reduced. Prevalence of tax evasion, which has similar effects to a subsidy, in Galapagos tourism and fisheries sectors is unknown.

Fisheries control, management and research continue to depend on tourism fee and donors. Fuel subsidies eliminated. Tax system tightened up.

Legislation is introduced to enable tariffs to be charged to fishermen, boat owners, and traders in fisheries products. Tariffs are introduced that cover part of the costs.

Each sector pays fees that at least cover costs of control, management and research. Subsidies, if any, are for actions to reduce impacts on the natural environment, for the public good.

CDF/WWF Biodiversity Vision Chapter 11 – The Vision and Key Issues

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CHAPTER 11 – SUMMARY OF THE VISION FOR CONSERVATION OF GALAPAGOS BIODIVERSITY AND THE ISSUES CENTRAL TO ITS ACHIEVEMENT

Principal Authors – R. Bensted-Smith, T. Allnutt, G.M. Branch, R.H. Bustamante, C. E. Causton, E. Dinerstein, G. Powell, H. Snell, A. Tye, G.M. Wellington and J.D. Witman

11.1 Summary of Marine Biodiversity Vision

The marine ecosystem of the Galapagos Marine Reserve is a diverse, climatically variable, insular marine environment with high levels of endemism. Unlike the terrestrial counterpart of the ecosystem, where the ecological changes caused by invasive species have acquired their own powerful momentum, most elements of the marine ecosystem could return to a natural state, if human activity in and around the eco-region were to be controlled. Because of this, the marine biodiversity vision is less limited than the terrestrial vision by problems of irreversible change that have already happened, or inevitable future losses, although some specific features, such as the abundance of cetaceans, may take much longer than 50 years to recover. By intensifying the El Niño phenomenon in Galapagos, global warming could have a substantial impact on the marine ecosystem, especially if it interacts with local anthropogenic impacts, such as over-fishing. The best insurance against major loss of biodiversity due to global warming is to maintain a healthy, robust ecosystem, not already stressed by over-exploitation. This is what we seek in the vision for marine biodiversity conservation 50 years hence, which has been elaborated in the tables of Chapter 10 and which responds to the conservation objectives and policies already adopted by Ecuador for the GMR and Galapagos as a whole. It can be summarized as follows:

• At least 36% of the coastal zones (including all flora and fauna directly influenced by the sea and extending out to two nautical miles) would be protected within No-Take Areas (NTAs) i.e. areas where no extraction of resources are allowed.

• In all biogeographic regions, all habitat types in both coastal and offshore waters would have at least 20% of their area protected within NTAs.

• The functioning and structure of biological communities would be maintained. Within the NTAs they would be restored to pre-exploitation condition. Natural levels of diversity of communities, species and genotypes would be maintained, as would trophic complexity, habitat structure, population connectivity, resilience to external changes, recruitment patterns and water quality. The introduction of pathogens or other alien species would be prevented.

• Sustainable populations of all species would be maintained. Within the NTAs they would be restored to pre-exploitation levels. Populations of herbivorous fish, seaweeds, gorgonians, urchins, hermatypic corals and marine iguanas would all be maintained, whilst the exploited populations of black corals, lobsters, sea cucumbers, conch, snapper, grouper and sharks would be restored.

• In areas of the GMR that fall outside of NTAs, resources would be managed to ensure sustainable use and human activities would be regulated to ensure that they have no significant negative impacts on biodiversity and evolutionary processes, and to allow recuperation of any degraded biological systems wherever possible. There would be no exploitation of species with key ecological roles, such as herbivorous fish, sea urchins, barnacles and predatory snails. Marine mammals would be fully protected. By-catch problems would be negligible.

• Species that occur in the GMR but range beyond its boundaries would be protected by (a) establishment of marine protected areas on mainland Ecuador, (b) special protection granted to individual species or groups of species, such as turtles, albatross, whales and sharks, and (c) international treaties and agreements.


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• The marine environment and biodiversity would be maintained and would continue to provide a high-value resource for tourism, fisheries and other human uses of the sea. Tourism and fisheries would become complementary, not antagonistic, operations.

• All utilisation of resources would be managed in such a way as to achieve conservation goals, whilst maximising the long-term benefits to the people of Galapagos.

Of these conditions the most critical is the long-term guarantee that biodiversity and ecological structure and functioning will be sustained in fully protected areas that cover at least 36% of the coastal areas and 20% of all habitats.

The ultimate aim is to have a healthy, thriving marine ecosystem, characterised by a diverse and abundant native flora and fauna with appropriate structure and function, which will allow future generations to use optimally and benefit from this rich heritage.

11.2 Summary of Terrestrial Biodiversity Vision

The vision for terrestrial biodiversity 50 years on is constrained by the fact that we must slow down, halt and eventually reverse the accelerating trend towards ecological degradation and biodiversity loss caused by alien species that have already been introduced. Already the number of introduced plant species exceeds the number of native plant species, whilst the rates of introduction of new plant and animal species, are thousands of times higher than estimated natural rates of establishment during the millions of years before people reached Galapagos. Nevertheless here, unlike other oceanic archipelagos, the long-term ambition of restoring almost all the original terrestrial biodiversity is still within the bounds of the possible, albeit not within the coming half century. A second threat to be dealt with is the fact that much of the humid highland habitat lies within agricultural zones and has already been degraded or lost. Only 7% and 26% of the humid areas on San Cristobal and Santa Cruz respectively lie within the National Park. The agricultural zones also serve as centers for alien species establishment and subsequent spread into the remaining fragments of natural habitat. A third problem, much more limited in scope, is that of unsustainable rates of harvesting, for example of trees, or killing, for example of birds by vehicles. The biodiversity vision, based on assumptions of optimal management to address these three problems and restore endangered species, can be summarised as follows.

• There would be no additional species extinctions of plants or vertebrates. Below species level, the additional extirpations would not exceed 1% of present numbers of subspecies and distinct populations respectively.

• The proportion of biodiversity (species, subspecies and populations) which is recognised under IUCN categories as being critically endangered or endangered would reduce from 24% (plants) and 50% (vertebrates) to 10% and 30% respectively. All extant species would have a distribution and abundance at least 50% of that prevailing in pre-human times and their within species genetic diversity would be on the increase. Within the Park this figure would rise to 80%.

• Of the 36 identified plant communities the number having all their distinct sub-communities represented in areas isolated from anthropogenic disturbances would rise from 8 to 32.

• The proportion of humid highland habitat that is uncleared and has a species composition not significantly altered from the pre-human baseline should be increased from the current one third (by area) to at least half. This would involve restoring natural habitat in the agricultural zones, in order to maintain viable natural biological communities, as well as combating invasive species throughout the inhabited islands. On business-as-usual trends the proportion is projected to halve to less than 20%, with virtual elimination of unmodified biological communites in San Cristobal, Santa Cruz and Floreana, and less than 10% remaining in southern Isabela.

• The approximately 11% of the archipelago’s area that lies outside the inhabited islands and Santiago, would be composed of near pristine islands. Within the inhabited islands the extreme fragmentation of highland habitats would be reduced by restoring habitat in some inter-linking areas.


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• The reduction in distribution and abundance of alien species should reduce substantially the number of species experiencing unnatural biotic interactions and selective pressures. Enhanced preventive measures, especially quarantine inspection, education and curbing the volume of transport, should reduce the introduction of alien species and the number of species experiencing increased gene flow between formerly isolated populations. Conversely, the reduced gene flow caused by habitat fragmentation should be mitigated by habitat restoration. Key elements of the vision with regard to alien species are that:

- The rate of introduction and establishment of new alien species would be reduced by 2 orders of magnitude;

- Of the alien species already introduced, 10% of plants and 80% of vertebrates would be eradicated from the archipelago, many others would be reduced in distribution and abundance, and the effects of the remaining alien species would be mitigated;

- There would be a net decrease, relative to current status, in the distribution of alien species in the archipelago.

Most of the features of the terrestrial biodiversity vision depend on success in addressing the problem of introduced alien species. Under the optimal management scenario, which underlies the terrestrial biodiversity vision, intense prevention and monitoring efforts would reduce the rate of successful colonizations by new alien species to less than one per year. On the other side of the equation, complete eradication from Galapagos would have been achieved for most of the already introduced vertebrates and a significant percentage of the introduced plants. Furthermore, the impacts of alien species upon the native flora and fauna would in all cases be mitigated. In short, in 2050 the balance would have been tilted in favour of mitigation and restoration and against invasion

11.3 Integration and Comparison of the Marine and Terrestrial Visions

The marine and terrestrial visions are fully compatible but are strikingly different in the problems identified. However, for the many coastal species there is in fact more synergy than revealed by the separate analyses. Under business-as-usual scenarios the combined pressure of over-exploitation in the sea and introduced alien species on land could push some native species or populations rapidly towards extirpation. It may also make them more vulnerable to the marked environmental fluctuations (El Niño – La Niña), which characterise Galapagos. On the positive side, success in tackling both over-exploitation and alien species will accelerate restoration.

When we look at the key factors that will determine success or failure in achieving the conservation vision, the marine and terrestrial analyses are quite different, although with one fundamental factor in common. The principal difference is that in the marine environment the trends towards loss of biodiversity can be reversed by quantitative and qualitative changes in certain human activities, principally fishing, whereas in the terrestrial environment the ecological degradation has a life of its own, a momentum driven by introduced alien species. Furthermore, to minimize the link between human presence and the introduction of still more alien species represents a tough social and technical challenge.

The fundamental common factor, affecting both marine and terrestrial projections, is population: the number, distribution, development decisions, lifestyles and behaviour of the province’s residents. For marine conservation, the choice of economic livelihood between extractive and non-extractive activities is critical, whilst terrestrial conservation depends more on the quantities of transport to and within the archipelago, and the willingness to adopt an “island lifestyle” that favours the control of alien species. The absolute size of the human population is a major contributing factor in both cases. Indeed, the General Regulation of the Special Law for Galapagos already recognizes that the Galapagos regional plan must include measures to stabilize human population, promote a lifestyle compatible with a strategy for ecological and socio-economic sustainability, and apply “total control” of alien species in inhabited as well as protected areas. Section 11.4 considers this crucial issue in more detail.


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One area needing careful balancing of marine and terrestrial biodiversity considerations is tourism. For all biodiversity, tourism has the advantages of providing a non-extractive livelihood for residents, linking local economic development to the existence of abundant flora and fauna, and financing most conservation work in the Islands. It has been and will continue to be the principal economic activity in the Islands. For marine biodiversity conservation, conversion from the extractive activity of fishing to non-extractive tourism is unequivocally advantageous. However, the analysis of terrestrial biodiversity conservation identifies the risk that tourism expansion will exacerbate the introduction of alien species and their dispersal between islands and within the larger islands. The existing strict controls on tourism and the conservation ethic of many tour operators reduce the risk, but cannot eliminate it.

Lastly, it should be remembered that the marine and terrestrial projections and visions may be affected by external factors, which those responsible for managing Galapagos can do little or nothing to influence. Of these the most significant is likely to be climate change. For marine conservation, global warming is likely to lead to increasing intensity of El Niño phenomena, which might change the projections to favour those species with high tolerance to El Niño and reduce or eliminate those which are vulnerable to such conditions. For example, the Galapagos penguin would be at risk. However, this is largely speculative and climate change is unlikely to change the principal requirements for biodiversity conservation. A fully functioning ecosystem with a high percentage of No-Take Areas will be more resilient and better able to adapt to climate change without major loss of biodiversity.

The impacts of climate change on the terrestrial biodiversity are more difficult to predict, because the El Niño – La Niña cycle affects the dynamics of competition between native and alien species. Experience suggests that El Niño conditions favour arrival, establishment and range expansion of alien species in Galapagos. Climate change would also have a direct effect on the distribution of native flora and fauna. Nevertheless, as in the marine case, climate change would probably affect the specific projections of biodiversity loss or recovery, rather than the principal requirements for effective conservation of terrestrial biodiversity.

11.4 Key Requirements for Progress towards the Biodiversity Vision

Chapter 1 of this document described the commitments made by the Ecuadorian Government to protect the biodiversity and evolutionary processes of the Galapagos Archipelago. The subsequent analysis has attempted to elaborate those commitments into a vision of what the flora and fauna of Galapagos could be like in the year 2050, if the Islands were managed optimally for conservation, thereby providing sustainable benefits to the people of Galapagos.

The vision provides a 50-year goal, to orient policy makers and planners, and a benchmark against which to measure progress over the coming decades. The vision cannot prescribe the specific measures to be taken towards achieving the conservation goal, because the measures depend on a complementary analysis of the social and economic situation and development aspirations in relation to the factors affecting biological conservation. Nevertheless, the vision can identify crucial issues, that must be addressed if the loss of biodiversity is to be reversed. An important process in which the comprehensive analysis of these issues should take place is the preparation of the Galapagos Regional Plan, which started in 2000. The Special Regulations for the Special Law for Galapagos, which deal with (1) fisheries, (2) tourism, (3) environmental control and (4) alien species, quarantine and agriculture, should also take full account of these issues1, as should the continuous process of policy development and investment planning by INGALA, GNPS, local councils, national ministries and funding agencies. The big issues, upon which the conciliation of conservation and development aspirations will depend, can be summarised as follows:

• No-Take Areas: Respect for No-Take Areas in the Marine Reserve, and their eventual expansion to recommended sizes.

• Management of the artisanal fishing fleet: fixed number of boat permits with closed access, incentives to reduce and redistribute effort, boat capacity, fishing methods, internal discipline, enforcement.

1 The special regulations have been in preparation since the enactment of the SLG in 1998 but so far none of the four has been finalised and published. This is a serious limitation for conservation.


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• Quarantine inspection: Full establishment of quarantine inspection and monitoring system, with infrastructure, equipment and professional staff.

• Special Regulations: Development and application of regulations on fisheries, tourism, environmental control and quarantine/agriculture/invasive species.

• Management of passenger traffic and cargo to the archipelago and between islands: limited ports of departure and entry, volume of traffic, inspection, permitted products, conditions of shipment, economic incentives to reduce traffic/importation.

• Management of tourism and within-island travel: balancing local development, marine conservation and the alien species problem; limiting infrastructure and other dispersal risks; linking tourism opportunities to reduction in fishing fleet.

• Management of the agricultural zone: control of alien pests; local production to reduce imports; economic viability; livestock entering park; restoration of natural habitat on agricultural zone land through purchase, stewardship grants and other incentives.

• Habitat protection and restoration: protection of Park boundaries; control of extractive use of terrestrial resources; restoration of degraded habitat in Park.

• Restoration of endangered species: prioritisation of endangered species conservation; safe ex situ populations as back-up; contingency plans for responding to conservation emergencies.

• Urban planning and management: control of alien pests; control of pets; location and planning of settlements; management of waste.

• Population growth: application of residency controls; stabilization or reduction through family planning and incentives to adjust immigration-emigration balance; plan distribution within archipelago to minimize impacts, especially on threatened or sensitive habitats; education.

• Education: educational reform, involving conservation institutions and private sector, to produce a well educated, trained population, committed to a distinct, conservationist island lifestyle.

• Motivation and capacity building for local participation in conservation: making conservation relevant to people’s lives; building partnerships with stakeholders.

• Use of economic incentives and disincentives: establishment of secure mechanisms to encourage donor financing of incentives with conservation objectives (e.g. buy back of fishing permits); application of the principle that the polluter (or person whose activities put the ecosystem at risk of damage) pays.

• Control of alien species: investment in management of alien species, including prevention, control, mitigation and eradication; systematic prioritisation of efforts; training.

• Terrestrial research and monitoring: investment in research and monitoring related to the terrestrial biodiversity vision, including new fields such as alien pathogens and parasites; baseline inventory of species and their distribution.

• Surveillance and enforcement: investment in the capacity to patrol, protect and manage the protected areas; strengthening of the judicial system.

• Marine research and monitoring: investment in research and monitoring related to the marine biodiversity vision, including ecosystem functioning, the effects of No-Take Areas and the population biology of exploited species.

The decisions and actions on these issues will determine whether or not development will become sustainable, according to the definition in the Special Law for Galapagos, and whether progress will be made towards the vision for Galapagos biodiversity in the year 2050. Whilst


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many of the legislative, policy and planning issues are still under discussion, the Government of Ecuador has already demonstrated its willingness to invest in conservation, including a US$ 18 million Global Environment Facility project on invasive species and a US$ 10 million Inter-American Development Bank loan for investment in marine management, quarantine and institutional strengthening.

The scale of the challenge presented by invasive alien species, and the links between alien species and a wide range of policies on socio-economic development and population, have to be recognized. To illustrate this, Table 11.1 lists a sequence of steps in the process of invasion by an alien species and, for each step, lists some measures that would reduce the risk of invasion. The logic behind the table is as follows. The rate of introduction and spread of alien species can be viewed as the product obtained by multiplying the volume of traffic by a series of probabilities representing the risk that an organism passes each barrier in turn, becoming eventually a threat to biodiversity. To reverse the current loss of biodiversity and make progress towards the biodiversity vision, this rate of introduction and dispersal of invasive alien species has to be reduced by a factor of the order of between 100 and 1000 from the 1999 rate. Consequently, it will require a concerted set of measures aimed at reducing most factors in the multiplication chain and preventing the increase of others.

Table 11.1 Steps in the invasion process and potential mitigating measures

FACTORS WHICH, MULTIPLIED TOGETHER, DETERMINE THE RATE OF ALIEN INVASIONS

RANGE OF POTENTIAL MANAGEMENT MEASURES TO BE ASSESSED IN A COMBINED BIOLOGICAL AND SOCIO-ECONOMIC ANALYSIS

Number of people (residents and tourists)

Control of tourism volume. Family planning. Use of economic incentives and disincentives to influence immigration/emigration balance. Planning of urban land use, economic activities, education, training and employment.

Volume of traffic per person (people and cargo traffic)

Increase self-sufficiency of province and of each island, especially in agricultural products. Use economic incentives and disincentives to encourage self-sufficiency and discourage importation and excessive travel. Substitute telecommunications for travel.

Probability that organism gets on board in a given trip

Effective quarantine inspection service on mainland, with comprehensive lists of permitted and prohibited products. System of authorized suppliers for some agricultural products. Appropriately located, clean infrastructure and facilities, designed for quarantine inspection purposes. Education to develop awareness and commitment to minimize accidental introductions. Regulation, information and enforcement to deter deliberate introductions.

Probability that the organism survives the journey

Strict bio-security standards for boats and planes. Fumigation.

Probability that the organism is not detected by inspectors at arrival point

Strengthen quarantine inspection service in the islands, in terms of facilities, human resources and funds. Concentrate inspection effort by minimizing number of entry points into the islands. Strictly regulate landings and shore excursions at sites without inspection facilities e.g. tourism sites, fishing camps, research and management camps, hunting trips.

Probability that the organism survives after arrival

Concentrate traffic on sites that present a hostile environment for most alien species. Keep ports and airports clean. Increase vigilance during El Niño.


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Probability that a small founder population is established

Design within-island transport networks and services to minimize risk of transporting the organism from a hostile to a favorable environment. In other ways, this factor is largely dependent on the effectiveness of other measures e.g. reducing the number of individual organisms in the arriving group reduces likelihood of establishment. Also absence of vectors, food plants, commensal species etc hinders establishment.

Probability that the small founder population is not rapidly detected and eradicated

Regular, intensive monitoring around ports of entry and high risk sites (visitor sites, garbage dumps etc). Reduce within-island transport networks that may disperse invader before it is detected. Training of farmers and community as a whole to detect alien species and cooperate in their eradication. Research to develop detection and eradication methods. Develop contingency plans. Establish regulations to control possession, cultivation/rearing, release, transport and trade of prohibited alien species and their products.

Probability that the organism is or becomes invasive, either immediately or after an extended lag phase.

This is partially innate character of the species in the Galapagos environment, not influenced by management measures. Again presence/absence of vectors etc may be crucial. Also, influx of new genetic diversity to an established population may produce invasiveness. Climate change may change invasiveness of some species.

Probability that the invading organism is not eradicated or adequately controlled

Research and develop eradication and control methods. Increase resources available for eradication and control. Prioritise research, eradication and control activities for maximum effect. Establish powers and procedures to control alien species outside protected areas, and obligations and incentives for landowners and local authorities to cooperate.

Probability that the invading organism is able to have a significant impact on biodiversity

This depends on the ecological relationships between the invader and the native organisms, and on the extent of dispersal around the archipelago. Impacts would be less if human population were concentrated on less vulnerable sites rather than on sites that are biologically sensitive and/or are the gateway to large, biologically important areas. Management measures to influence human population distribution towards less vulnerable islands/areas and away from highly sensitive ones should be both regulatory and economic i.e. economic planning, investment policies, incentives and disincentives, and provision of better social services in. Design of inter-island transport system, tour-boat itineraries, control of fishing boat landings, road networks, provincial social services and other infrastructure are all important factors in dispersal and also affect human population distribution.

Probability that the impact on biodiversity is not mitigated by measures other than control

Research on native species ecology and conservation. Protection in situ of vulnerable species and habitats. Maintain breeding and repatriation programs. Ex situ conservation of endangered species. Prioritise mitigation efforts.


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The table presents an extensive array of potential measures, in order to highlight for decision-makers the wide range of decisions and actions in Galapagos that will determine, one way or another, the future of the archipelago’s ecosystems and biodiversity. Several of the measures are already being implemented by the Ecuadorian authorities, others could easily be adopted, and others would face strong social, economic or political objections. The next step in the planning process should be a combined socio-economic and biological analysis of all these potential measures, and others that may be developed in the course of the analysis, in order to come up with the optimal combination of measures, that can achieve the conservation objectives in a way compatible with principles of human welfare and social justice. It is worth noting that the term “social justice” implies fairness towards future generations as well as towards current vi sitors and residents of the islands.

Amongst the most difficult and sensitive measures to deal with will be those concerning human population and transport. Terrestrial biologists at the workshop concluded that, in order to achieve a balance between the rate of introduction of alien species and the capacity to deal with them, it would be necessary to reduce total human population and volume of transport (cargo, tourism, local travel etc). Many went further and considered that the alien species problem could be made manageable – and the biodiversity vision achievable – only if the number of sites of introduction and dispersal were to be limited by reducing the number of ports of entry into Galapagos and having, eventually, no more than two human population centers (in Santa Cruz and San Cristobal Islands). Could such radical social changes ever be feasible? Family planning and reproductive health services would generally be welcomed. Relocation schemes could be totally voluntary, providing incentives and assistance for those who choose to move to another island or to renounce their permanent resident status and move to the mainland, and thereby responding to the desires of those who apply for such assistance. Such population changes could also bring short-term benefits to remaining residents, in terms of space for housing, access to social services and education, share of benefits from natural resources and so on, in addition to the obvious long-term benefit of preserving the natural resource base of the local economy. Nevertheless, it is evident that the social aspects of the alien species problem need profound analysis and discussion with all concerned, as well as careful handling, whilst the socio-economic incentives would have to be highly creative, given the big socio-economic disparity between Galapagos and the mainland.

The political aspects of population issues are difficult anywhere, but especially so in a society noted for conflicts between visionaries, focused on the long-term future, and populists, focused on short-term ambitions. The tendency to conflict is perhaps not surprising, given the high proportion of recent economic migrants and the plentiful opportunities for misunderstanding – or even distorting - the complex subject of sustainability in island ecosystems. On the other hand, Galapagos has been a pioneer in Ecuador in promoting public discussion of conservation and development issues and establishing formal structures for local participation in decision-making. Creative, open-minded thinking will be needed!

So will it be possible for Ecuador to design and implement a socially, economically and politically feasible combination of measures, which will achieve a 100- to 1000-fold reduction in invasions by alien species? On the answer to this question depends the hope for long-term conservation of the terrestrial biodiversity of Galapagos. It is the toughest challenge facing the Galapagos regional planning process and the Government of Ecuador, as it seeks to develop a comprehensive strategy to harmonize biodiversity conservation requirements with social and economic aspirations. A biodiversity vision has been articulated. It should be complemented by social and economic analysis, in order to produce a comprehensive biological and social vision and to plan the optimal combination of management measures.

11.5 Monitoring Progress Towards the Vision

For the biodiversity vision to serve as a benchmark for measuring progress over the coming 50 years, it is necessary to monitor the indicators used to define the vision. Chapters 4 and 8 and their respective annexes describe the indicators. To measure all the indicators thoroughly and regularly would be a very expensive commitment, so CDRS scientists, with support from WWF and Fundación Natura, are working on an ecological monitoring plan, aimed at collecting and analyzing priority data, with sufficient sampling to be statistically robust and to distinguish anthropogenic from natural fluctuations (H.Snell et al. 2001). The CDRS has also been


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developing its use of Geographic Information Systems and Global Positioning System technology, which is revolutionizing the collection and analysis of field data and opening up new possibilities in ecological monitoring in Galapagos.

The draft monitoring plan focuses on terrestrial and coastal species. In the case of the marine environment, a number of key indicators are presently being monitored as part of the Management Plan for the Galapagos Marine Reserve. The Management Plan requires the CDRS to monitor marine biodiversity and populations of key indicator species of fishery and touristic importance within different Marine Reserve zones over the coming four years, in order to assess after two and four years the initial effects of the provisional zoning scheme. In these short periods only certain kinds of change can be detected; many important benefits of No-Take Areas take much longer to materialize. Nevertheless, it is important to predict and measure such changes as may occur, in order to provide technical input to the intended review and potential modification of the zoning scheme at those intervals. Many other factors, in addition to ecological data, will be presented for consideration during the review of the zonation system.

Financing will have to be sought for implementation of the terrestrial and marine ecological monitoring plans, since both demand a great deal of scientist time for field work and analysis, as well as data processing equipment.

REFERENCES

H. Snell, C.E. Causton and A. Tye (2001) “Ecological Monitoring for the Galapagos Archipelago: a Proactive Program for the Conservation of Biological Diversity” Charles Darwin Foundation, Galapagos Islands.

CDF/WWF Biodiversity Vision Annex 1.1 Biodiversity Workshop Participants

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ANNEX 1.1 – LIST OF PARTICIPANTS At the Biological Assessment Workshop Held in Galapagos, 25-28 May, 1999

Last/First Name

Country / location E-mail

Adsersen, Henning Denmark Copenhagen

[email protected]

Allnutt, Tom USA Washington

[email protected]

Anderson, Dave USA Wake Forest

[email protected]

Bensted-Smith, Robert Galapagos [email protected] Bostford, Loo W. USA

California [email protected]

Branch, George South Africa Cape Town

[email protected]

Bustamante, Rodrigo Galapagos / Australia

[email protected]

Causton, Charlotte Galapagos

[email protected]

Cifuentes, Miguel Costa Rica Turrialba

[email protected]

Cowie, Rob USA Hawai

[email protected]

Cruz, Eliecer Galapagos [email protected] Christensen, Villy Canada

Vancouver [email protected] [email protected]

DeVries, Tjitte

Ecuador, Quito

[email protected]

Dexter, Nick

Australia [email protected]

Dinerstein, Eric USA, Washington

[email protected]

Dowler, Robert USA, Texas

[email protected]

Eldredge, Lucius G.

USA, Hawaii

[email protected]

Espinoza, Fernando Ecuador, Quito

[email protected]

Ford, Glenn USA,

[email protected]

Gardener, Mark

Australia [email protected] [email protected]

Gaybor, Nikita

Ecuador, Guayaquil

[email protected]

Geller, Jonathan B. USA California

[email protected]

Gibbs, James

USA, New York

[email protected]

Godbey, Maria USA, Washington

[email protected]

Hickman, Cleveland

USA, Virginia

[email protected] [email protected]

Kakabadse, Yolanda Ecuador, Quito

[email protected]

Kendrick, Gary Australia [email protected] Marquet, Pablo Chile

Santiago [email protected]

CDF/WWF Biodiversity Vision Annex 1.1 Biodiversity Workshop Participants

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Martinez, Priscilla Galapagos / Australia


McCosker, John USA California

[email protected]

Merlen, Godfrey Galapagos [email protected] Mooney, Harold USA

California [email protected]

Muñoz, Edgar Galapagos [email protected] Nafus, D.M.

USA Idaho

[email protected]

Paulay, Gustav Guam [email protected] Peck, Stewart Canada

Vancouver [email protected]

Pellerano, Miguel

Argentina Buenos Aires

[email protected]

Porter, Sanford USA Florida

[email protected]

Powell, George

USA Washington

[email protected]

Reck, Gunter Ecuador Quito

[email protected]

Rejmanek, Marcel

USA California

[email protected]

Richmond, Robert Guam [email protected] Rivera, Fernando

Galapagos / Australia


Ruiz, Ruth Elena

Ecuador, Quito

[email protected]

Seddon, Mary

England, London

[email protected]

Silva, Paul

USA, California

[email protected]

Smith, Cliff

USA Hawaii

[email protected]

Snell, Howard

Galapagos / USA (New Mexico)

[email protected]

Spurrier, Lauren

USA, Washington

[email protected]

Suarez, Luis

Ecuador, Quito


Tye, Alan Galapagos [email protected] Ulloa, Robert

Ecuador, Quito

[email protected]

Valle, Carlos

Ecuador, Quito

[email protected]

Vargas, Hernán Galapagos [email protected] [email protected]

Veitch, Dick

New Zealand [email protected]

Wellington, Jerry

USA, Texas

[email protected]

Wikelski, Martin

USA Princeton

[email protected]

Witman, Jon

USA Virginia

[email protected]

CDF/WWF Biodiversity Vision Annex 1.2 Laws and Policies

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ANNEX 1.2 –Laws And Policies Concerning The Ecuadorian Government’s Conservation Goals For Galapagos

1. Introduction

The Ecuador Government has stated its objectives and policies for the conservation of the Galapagos archipelago in a hierarchy of documents, ranging from international instruments and the national constitution to policy papers and management plans.

This Annex provides an overview of the legal framework then compiles pertinent extracts from official Ecuadorian documents. The extracts highlight policies, objectives and regional approaches to planning. Also included are a number of articles illustrating the recognition of invasive alien species as the principal threat to terrestrial biodiversity and requiring that it be dealt with. The Special Regulations for putting this into practice are still in preparation. A balanced understanding of the laws, including their social, institutional and economic aspects, would require a more complete compilation than that presented here.

2. Overview of the Legal Framework

As a natural protected area, the Galapagos National Park is covered by the Forestry Law (1981) Amongst other functions ascribed by the law to the responsible ministry are to protect and prevent elimination of species of flora and wildlife, that are threatened or in process of extinction, and to fulfill and enforce fulfillment of national and international agreements. According to the General Regulations under that law, the objectives of the national protected area system include the preservation of outstanding resources, founded on ecological principles, and the perpetuation in a natural state of representative samples of biotic communities.

The particular activities to be allowed in a protected area depend on the management category. The official Procedures Manual for protected areas describes the objective of a national park as being the maintenance of the area in its natural condition, with any kind of exploitation or settlement being prohibited. In the case of the Galapagos National Park, according to the most recent management plan (published in 1996), the fundamental objective is to protect natural areas and scenic beauty of national and international significance, for scientific, educational and recreational use. Management attempts to maintain the biodiversity and the unique natural resources of the islands in the most natural state possible, but exploitation of resources is permitted in some zones.

These policies and objectives for the Galapagos National Park were largely unaltered by the enactment in 1998 of the Law for a Special Regime for the Conservation and Sustainable Development of the Province of Galapagos. Known as the Special Law for Galapagos (SLG), this legislation is highly significant for the orientation and integration of conservation and development in the province as a whole – protected and inhabited, marine and terrestrial – but does not modify the management regime for the park itself, except for one dangerous article, which in a list of the resources of the National Institute for Galapagos (INGALA) includes 2% of the surface area of the inhabited islands. The article has internal contradictions: the area delimited is supposed to be a “resource” of INGALA, which would appear to give INGALA property rights, but the use of the area has to be subject to the Park management plans as well as INGALA Council decisions, which would appear to give INGALA only use rights. Hopefully, this issue can be legally resolved in favour of maintaining the integrity of the Park.

An important issue for the near future is that new legislation concerning biodiversity and protected areas is in the pipeline, superseding the Forestry Law. A technical group has been drafting the biodiversity legislation, but it is unclear when this may come to Congress, because of other legislative priorities. The implications for the Galapagos National Park are not yet known but it is unlikely that the general objectives and principles for management of the Park will change.

For the marine component of the Galapagos ecoregion, the SLG brought dramatic changes in conservation status and objectives. Since 1986 the waters around Galapagos, for a distance of


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15 miles, had been a “Marine Resources Reserve” and in 1992 a well prepared management plan for the Reserve was published. However, neither industrial nor artisanal fishing sector respected the protected status, the shared jurisdiction between the Ministry responsible for Fisheries, the Navy and the National Park Service did not function effectively, and the legal status was weak, since the category of Marine Resources Reserve did not exist in the national protected areas legislation. After an intense national political debate, the SLG replaced this inadequate protection with a new legal and institutional framework, based on a consensus reached in the Archipelago between the Park, the Charles Darwin Foundation and local stakeholders in the marine environment, including fishermen, tour operators and naturalist guides.

The SLG creates a new category of national protected area, the Marine Reserve, which aims to guarantee the protection and maintenance of the biodiversity in the long term, and at the same time provide a sustainable flow of products, services and uses for the benefit of the community. Amongst the requirements in the legal description is that there must be participatory management. Under this category the SLG establishes the Galapagos Marine Reserve (GMR), including the internal waters of the archipelago and the surrounding waters to a distance of 40 miles. Thus, the GMR has the legal protection afforded by the Forestry Law to all protected areas and also by the specific provisions of the SLG; the latter prohibits industrial fishing and includes stiff sanctions against illegal fishing of any kind.

Unfortunately, the fisheries regulations, and all the Special Regulations of the SLG, which were due in June 1998, have yet to be produced. This prevents full implementation of the law and calls into question the commitment to its conservation provisions. Much work was done in 2001, under the auspices of the Ministry of Environment, to finalise the Special Regulations on Fisheries and on Tourism, but in 2002 it seems that momentum has been lost. The Special Regulation on Quarantine, Agriculture and Introduced Species is also well advanced, whilst the Special Regulation on Environmental Control, where environmental impact assessment procedures are defined, will take a few more months.

A framework Management Plan for the GMR, based on a local consensus achieved through a lengthy participatory process, was approved in March 1998. In the Plan, the general goal for the GMR is to protect and conserve the archipelago’s marine and coastal ecosystems and their biological diversity for the benefit of humanity, local populations, science, and education. The Plan lists a dozen specific objectives, the majority oriented towards conservation, plus others about the main uses i.e. tourism, fishing and scientific research. Despite opposition from the industrial fishing sector and the difficulties resulting from the lack of regulations, the SLG is in effect and provides a good legal and institutional framework for the conservation of the GMR.

The SLG seeks to harmonize conservation and sustainable development in the archipelago, by establishing principles, rules, responsibilities and privileges applicable throughout the province, thereby responding to conservationists’ insistence that the Park cannot be preserved if the problem of invasive species is addressed only within the Park, whilst the root causes of the problem and the points of entry for the invaders both lie within the inhabited areas. Since restricting rights of residence was one of the key requirements, the Constitution of Ecuador had to be changed, in order to enable a province-specific legislative regime to be established. The SLG ranges over many sectors, reflecting the participation of many interest groups in its preparation, but lacks a clear strategy for achieving the ambitious goal of harmonising conservation and development. Nevertheless, the law establishes important policies and principles, starting with the preliminary considerations, which recognize the ecological interconnections between the Park, Marine Reserve and inhabited areas, identify introduced species as the principal threat, and declare that it is State policy to protect and conserve the terrestrial and marine ecosystems of the province of Galapagos, its exceptional biological diversity and the integrity and functionality of particular ecological and evolutionary processes for the benefit of humanity, the local population, science, and education. The same issues are addressed in a list of seven principles at the start of the law. In the same lists of considerations and principles appear issues of human development and welfare. The glossary of the law includes a definition of sustainable development, which requires that the development activity avoid exacerbating the introduced species problem. Thus, it is clear that the success of the law in conserving biodiversity will depend both on its correct application and on finding a strategy and practical mechanisms by which conservation and development can indeed be harmonised. The document in which the strategy and mechanisms will be defined is the Regional Plan. Guidelines for this plan, which is binding on all authorities in Galapagos, are included in the


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General Regulation for the application of the SLG, which was promulgated in January 2000. The Regional Planning process started in 2000 and should be completed in the course of 2002.

Since the SLG was enacted, a national Environmental Management Law has also been enacted (July 1999). This brief law provides a framework for a decentralized, participatory system of environmental management, leaving much of the detail to future regulations. For Galapagos it adds little to the environmental protection afforded by the SLG but provides useful guidance for the environmental impact assessment and environmental audit components of the Special Regulation on Environmental Control, to be prepared under the SLG.

This plethora of laws, plans and policies all reaffirm the intention of the Government of Ecuador to conserve the biodiversity and ecological and evolutionary processes of Galapagos, whilst promoting compatible local development. This commitment is made international by the status of Galapagos as both a UNESCO World Heritage site (extended in December 2001 to include the Galapagos Marine Reserve) and a UNESCO Man and Biosphere Reserve. Thus, the elaboration by scientists of a biodiversity vision can help Ecuador articulate in concrete, specific terms the conservation goal, which it has set itself.

3. Excerpts from Laws, Policies and Plans

Excerpt 3.1

LAW OF FORESTRY AND OF NATURAL AREAS AND WILDLIFE

Ley No. 74

Registro Oficial No. 64 of 24 August 1981

SECTION II

ON NATURAL AREAS AND FLORA AND WILDLIFE

CHAPTER I

On the National Heritage of Natural Areas

Art. 71. The heritage of natural areas of the State must be conserved unaltered. To this end plans for regulating each one of said areas will be formulated.

This heritage is inalienable and not subject to termination (“imprescriptible” in Spanish), and no real (property) rights can be established over them.

. . .

Art. 76. The flora and wildlife are under the dominion of the State and their conservation, protection, and administration corresponds to the Ministry of Agriculture and Livestock, for which it exercises the following functions:

a) Control hunting, collection, apprehension, transport, and traffic of animals and other elements of the wild flora and fauna;

b) Prevent and control the contamination of the soil and waters, as well as the degradation of the environment;

c) Protect and prevent the elimination of the species of wild flora and fauna that are threatened or in the process of extinction;

d) Establish breeding centers, nurseries, wild plant gardens, and research stations for the reproduction and protection of the wild flora and fauna;


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e) Develop activities that demonstrate the domestic use and advantage of the flora and wildlife, by means of methods that avoid diminishing their integrity;

f) Fulfill and enforce fulfillment of national and international agreements for the conservation of the wild flora and fauna and their environment; and

g) Others assigned by the Law and Regulations.

Excerpt 3.2

GENERAL REGULATIONS OF APPLICATION TO FORESTRY AND THE CONSERVATION OF NATURAL AREAS AND WILDLIFE

Executive Decree No. 1529

Registro Oficial No. 436 of 22 February 1983

SECTION II - ON NATURAL AREAS AND FLORA AND WILDLIFE

CHAPTER I

On Natural Areas

Art. 197 The establishment of the system of natural areas of the State and the management of the flora and wildlife are governed by the following basic objectives:

a) To tend toward the conservation of renewable natural resources in accordance with social, economic, and cultural interests of the country;

b) To preserve the outstanding resources of flora and wildlife, scenery, historial and archaeological relics, founded on ecological principles;

c) To perpetuate in a natural state representative samples of biotic communities, physiographic regions, biogeographic units, aquatic systems, genetic resources, and wild species in danger of extinction;

d) Provide opportunities for the integration of humans with nature; and

e) Assure the conservation and protection of wild flora and fauna for its rational use to benefit the population.

Art. 199. The activities allowed in the System of Natural Areas of the State are the following: preservation, protection, investigation, recuperation and restoration, education and culture, controlled recreation and tourism, controlled sport fishing and hunting, rational use of the wild flora and fauna.

These activities will be authorized by the National Forestry Department, based on the management category of the natural areas. (Article substituted by Decreto Ejecutivo No. 857, Registro Oficial No. 213 of 24 June 1985).

Excerpt 3.3

PROCEDURES MANUAL: OBJECTIVE OF A NATIONAL PARK

Maintenance of the area in its natural condition, for the preservation of the ecological, aesthetic and cultural features, with any kind of exploitation or settlement being prohibited.


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Excerpt 3.4

MANAGEMENT PLAN OF THE GALAPAGOS NATIONAL PARK (1996)

CHAPTER 4: POLICIES AND PRINCIPLES

The Galapagos National Park (GNP) attempts to maintain the biodiversity and the unique natural resources of the islands in the most natural state possible. In accordance with the universally accepted definition for this category of protected area management, its fundamental objectives are to protect natural areas and scenic beauty of national and international significance, for scientific, educational, and recreational use.

The GNP supports development in the insular region, in agreement with the characteristics and capacities of the unique Galapagos ecosystems. On several occasions, policies and procedures have been defined and ratified, to support selective and special development in accordance with the particular conditions of the islands.

For more than three decades, the Ecuadorian Government has expressed its willingness to conserve the natural resources of the Archipelago through concrete administrative and management actions.

All the management decisions that affect the GNP are regulated by the general outlines expressed in the Management Plan. These decisions must be taken on the basis of appropriate technical and scientific information. The implementation of the Management Plan is carried out through annual operation plans and specific thematic plans.

Excerpt 3.5

MANAGEMENT PLAN FOR CONSERVATION AND SUSTAINABLE USE FOR THE GALAPAGOS MARINE RESERVE (1999)

2. GOAL AND OBJECTIVES

2.1. GENERAL GOAL

The general management goal for the Galapagos Marine Reserve is:

“To protect and conserve the Archipelago’s marine-coastal ecosystems and their biological diversity for the benefit of humanity, local populations, science, and education.”

2.2. SPECIFIC OBJECTIVES

The specific management objectives that derive from the general goal are:

a) Protect and conserve the Galapagos marine and coastal ecosystems to maintain evolutionary and ecological processes in the long term.

b) Complement, with the marine and coastal components of the Galapagos ecosystems, the protection of terrestrial environments and of communities and species of protected flora and fauna that depend on the marine environment for their survival.

c) Protect the marine and coastal species that are important due to being endemic, vulnerable, and for their genetic, ecological, touristic, or intrinsic values.

d) Ensure the maintenance and preservation, or in certain cases, the recuperation of populations of species of fisheries resources that have great commercial importance for the fishery.


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e) Facilitate that Galapagos fishermen maintain and improve their social and economic base, ensuring that the realization of fishery activities are compatible with the biodiversity.

f) Conserve the marine and coastal ecosystems of Galapagos as the economic base of the activity of tourism, by controlling, preventing, and mitigating the environmental impacts caused by this activity.

g) Provide and promote scientific research to increase knowledge on marine biodiversity, on exploited sites and species, and on the ecological impacts caused by human activities.

h) Provide and promote scientific and cultural education on marine and coastal nature.

i) Implement and practice a system of adaptive and participatory management of the Marine Reserve, through which follow-up data may be used to modify the management according to new information or to socioeconomic and environmental issues.

j) Create and strengthen both structures and permanent funding through the GNP to implement the work of the Participatory Management Council of the Marine Reserve.

k) Ensure the preservation and maintenance of the scenic values present in the marine and coastal ecosystems.

l) Establish the basic scientific and technical requirements to ensure environmental protection and the conservation and sustainable development of natural resources in the Marine Reserve.

Excerpt 3.6

CONSTITUTION OF THE REPUBLIC OF ECUADOR (August 1998)

CHAPTER IV. ON SPECIAL REGIMES

Art. 238 [Special Regimes for Territorial Administration]

Special regimes will exist for territorial administration, on the basis of demographic and environmental considerations. For the protection of the areas subject to a special regime, it will be possible to limit within them rights of internal migration, work or any other activity that may affect the environment. The law will regulate each special regime.

The residents of the area in question, affected by the limitation of their constitutional rights, will be compensated through preferential access to the benefits of the available natural resources and to the membership of associations which assure the heritage and family welfare1. In other matters, each sector will be governed according to what is established in the Constitution and the law.

. . .

1 Whilst the concept of benefits to compensate restricted rights may be appropriate in the case of indigenous tribes in the Amazon rainforest, it seems inappropriate to the Galapagos Islands, where the standard of living far exceeds the mainland, a high proportion of the population are recent economic migrants, few people can claim Galapagos origins back more than one generation, and the presence of humans has a negative impact on the national heritage, especially because of the unavoidable link to invasive species. It would have been more appropriate to state that to live in Galapagos is a privilege, which brings with it responsibilities to minimize one’s impact and contribute to conservation. The Special Law for Galapagos would still have included privileges and preferential treatment for residents, but these would have been privileges, that must be justified on the grounds of contributing to conservation, rather than constitutional rights.


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Art 239 [Special Regime for Galapagos. National Galapagos Institute.]

The National Galapagos Institute, or whoever may fulfill its responsibilities, will carry out the provincial planning, will approve the budgets of local dependencies of the national government and autonomous local authorities, and will oversee the execution of the budgets. It will be directed by a council composed of the governor, who will preside over it; the mayors, the provincial prefect, representatives of scientific and technical areas, and other persons and institutions whom the law may determine.

The provincial planning carried out by the National Galapagos Institute, which will have technical and scientific assistance and the participation of local dependencies of the national government and autonomous local authorities, will be unique and binding.2

Excerpt 3.7

LAW FOR A SPECIAL REGIME FOR THE CONSERVATION AND SUSTAINABLE DEVELOPMENT OF THE PROVINCE OF GALAPAGOS (18 March 1998)

Registro Oficial No. 278, Ley No. 67

In consideration:

That it is the duty of the State of Ecuador to oversee the conservation of the National Heritage of Natural, Terrestrial, and Marine Areas, as well as the development of surrounding human settlements, and to adopt legal measures established to provide a harmonious relationship with the inhabitants settled in the province of Galapagos;

. . .

That it is the policy of the State of Ecuador to protect and conserve the terrestrial and marine ecosystems of the province of Galapagos, its exceptional biological diversity and the integrity and functionality of particular ecological and evolutionary processes for the benefit of humanity, the local population, science, and education;

PRELIMINARY SECTION

Art. 2. BASIC NORMS FOR THE ESTABLISHMENT OF POLICIES AND THE PLANNING OF THE PROVINCE OF GALAPAGOS

The activities to establish the policies, planning, and execution of public and private works in the province of Galapagos and the area constituting the Galapagos Marine Reserve will be governed by:

1. The maintenance of ecological systems and the biodiversity of the province of Galapagos, especially native and endemic, allowing at the same time the continuation of the processes of evolution of these systems under minimal human interference, taking into account particularly the genetic isolation between the islands and between the islands and the continent;

2. Sustainable and controlled development within the carrying capacity of the ecosystems of the province of Galapagos;

3. Privileged participation of the local community in development activities and in sustainable economic use of the ecosystems of the islands, based on the incorporation of special models of production, education, training and employment;

2 The binding nature of the provincial (i.e. regional) plan is crucially important in enabling integrated management of Galapagos. On the other hand, the stipulations on composition of the INGALA Council, superimposed on prior stipulations in the SLG, have produced an unfavourable imbalance between short-term political and economic interests and long term conservation and sustainability interests.


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4. The reduction of the risk of introducting diseases, pests, and species of plants and animals that are exogenous to the province of Galapagos;

5. The quality of life of the resident of the Province of Galapagos must correspondto the exceptional characteristics of the World Heritage site.

6. The recognition of the interactions existing between the inhabited zones and the protected terrestrial and marine areas, and therefore the need for their integrated management;

7. The application of the precautionary principle in the execution of works and activities which could have effects contrary to requirements for protection of the environment or the insular ecosystems.

The regulatory instruments derived from this Law will include the scientific and technical requirements that ensure the protection of the environment, the conservation of natural resources and sustainable development

SECTION I - INSTITUTIONAL FRAMEWORK

CHAPTER III. ON THE TECHNICAL SECRETARIAT OF INGALA

Art. 10. RESOURCES OF INGALA

The resources of INGALA are composed of:

. . .

7. Two percent of the surface of the inhabited islands, which will be delimited by the Directorship of the Galapagos National Park. The delimitation and use of said areas will be subject to the corresponding management plans made by the Galapagos National Park, to the principles and norms established in this Law, and to the policies and decisions of the INGALA Council.

CHAPTER 1V, PARAGRAPH 1. ON THE MARINE RESERVE OF THE PROVINCE OF GALAPAGOS

Art 12. The Marine Reserve of the province of Galapagos is added to the category of Marine Reserve, of multiple use and integrated administration, according to the classification found in the title of the legal reforms, which is part of this Law (see Art 72 below).

The Marine Reserve comprises the marine zone within forty nautical miles of the baseline enclosing the Islands and the interior waters, according to the Executive Decree No. 959-A of June 28 of 1979, Official Register No.265 of July 13 of 1971.

Art 16. A minimum protection area is established out to 60 nautical miles from the baseline, to regulate the transport of toxic or high-risk products in this zone. These limits may be increased in conformity with international agreements and scientific research that may be undertaken for this purpose.

SECTION II – ON THE REGIME FOR RESIDENCE IN THE PROVINCE OF GALAPAGOS

Art 24. GENERAL PRINCIPLE

Everyone who enters or stays in the province of Galapagos must legalise their migratory status, in accordance with this Law, the General Administrative Regulations for applicaiton of this law and the Special Administrative Regulations on the subject.

The INGALA Council’s Committee for the Evaluation and Control of Residence, in accordance with this Law and its Administrative Regulations, will execute the control of residence.


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SECTION IV - PRODUCTIVE ACTIVITIES IN THE PROVINCE OF GALAPAGOS

CHAPTER 1. ON FISHERIES ACTIVITIES

Art 39. PRINCIPLES FOR FISHERIES ACTIVITIES

Fisheries in the Marine Reserve of the province of Galapagos will be subject to the principles of conservation, adaptive management and the guidelines for sustainable utilisation of hydrobiological resources, contained in this Law and the corresponding Management Plan.

CHAPTER III. ON AGRICULTURAL ACTIVITIES

Art. 53. POLICY ON AGRICULTURAL DEVELOPMENT

The agricultural activities in the province of Galapagos will be subject to the following criteria:

. . .

4. It is a duty of all people and legal entities to contribute towards the total control of introduced species and to the prevention of their entry and dispersion. The actions of inspection and quarantine will be given priority, as well as the total control and eradication of such species with aggressive behavior that affects the survival of native and endemic species of the Islands;

Art.54 TOTAL CONTROL OF INTRODUCED SPECIES

The Ministry of Agriculture and Cattle Farming, through the Ecuadorian Agricultural Health Service, SESA, will be in charge of carrying out inspection and quarantine actions, both of people and cargo, in ports and airports of departure and arrival, in coordination with the entities named in article 3 of the Special Administrative Regulations on Health and Quarantine for Agriculture and Natural Areas in the province of Galapagos. (This refers to a pre-existing regulation of the Ministry of Agriculture.)

Art.55 ERADICATION OF INVASIVE SPECIES IN AGRICULTURAL AREAS

The Annual Program for the Eradication of Animal and Plant Exotic Species, both in agricultural areas and sectors of the Galapagos National Park, will be prepared by the Ministry of Agriculture and Cattle Farming and the Forestry and Natural Areas and Wild Life Ecuadorian Institute (INEFAN). This will also be done in coordination with INGALA, the Charles Darwin Foundation and the legally recognized producer associations, under the principles of participatory management, which will also establish the corresponding obligations for executing the plan.

The financing for the execution of the annual program will come from the Inspection and Quarantine System and the Galapagos National Park.

Art.56 SPECIES PERMITTED TO ENTER GALAPAGOS

The Ministry of Agriculture and Cattle Farming and the Directorate of the Galapagos National Park, with the advice of the Charles Darwin Foundation and the research sectors of the province of Galapagos, will establish the regulations and procedures of total control (see definition in glossary below), inspection and quarantine and for approval for the introduction of any species.

SECTION V – ENVIRONMENTAL CONTROL

Art 61.

. . .


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Where necessary in accordance with the relevant regulations, prior to signing a public contract or to the issue of an administrative authorisation for the execution of public, private or mixed works, an evaluation of environmental impact will be required. The obligations arising from the evaluation of environmental impact will form part of said legal documents.

Art 62. PROHIBITIONS

The following are expressly prohibited:

....

5. The introduction of exogenous organisms to the islands, in conformity with the regulations in force;

6. The transportation, by whatever means, of animals, including domestic animals, from the mainland to the Islands, and of any indigenous species of fauna or flora or geological material from the Islands to the mainland or abroad; and,

7. The transportation between islands of indigenous or introduced organisms, without the corresponding authorisation.

SECTION VIII - REFORMS AND REPEALS

Art. 72. The following legal instruments are reformed or repealed:

a) The Forestry and Natural Areas and Wildlife Law

At the end of Article 109 of the Law of Forestry and of Natural Areas and Wildlife, the following paragraph must be added:

“Marine Reserve. -

Within the National Heritage of Natural Protected Areas, the category of Marine Reserve is created. The Marine Reserve is a marine area that includes the water column, the sea floor, and the subsoil, that predominantly contain unmodified natural systems, which are the object of activities of management to guarantee the protection and maintenance of the biological diversity in the long term, and at the same time to provide a sustainable flow of natural products, services, and uses for the benefit of the community.

. . .

SECTION IX

Art. 73. GLOSSARY

SUSTAINABLE DEVELOPMENT

There are three specific requirements for sustainable development in the province of Galapagos:

1. Biodiversity is maintained;

2. Evolutionary processes are maintained; and

3. There is no risk of causing, directly or indirectly, the introduction or dispersion of exotic species.

BIODIVERSITY

This term denominates the ecological diversity and the diversity of native and endemic biological species with all their variety of subspecies, races, geographically distinct populations, and genetic diversity in general.


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TOTAL CONTROL OF INTRODUCED SPECIES

By total control of species the following set of activities are understood:

1. To prevent the introduction to the Province of Galapagos of whichever species, genetic variety or modification of the flora or fauna, including micro organisms, which are not indigenous to Galapagos, except in the case of having specific authorisation as established in this Law;

2. To prevent the dispersal of species, varieties or modified forms in the Province of Galapagos, except with specific authorisation as established in this Law;

3. To prevent human interference in the distribution within the archipelago of indigenous species and of the genetic variety within each species;

4. To detect and eradicate new introductions to the Province of Galapagos and colonizations of new areas by exotic species already introduced;

5. To prevent the possession, cultivation, breeding or spreading in the environment of exotic species, except those allowed within the Administrative Regulations;

6. To eradicate the species already introduced, except those which are permitted by the Administrative Regulations; and,

7. To educate and train the inhabitants of the province of Galapagos to participate in control activities.

The restrictions described above apply to whole organisms and to any part of the organism capable of reproduction, including eggs, seeds, crops in vitro, cuttings, tissue or live samples of whatever type.

Excerpt 3.8

GENERAL REGULATION FOR APPLICATION OF THE SPECIAL LAW FOR THE CONSERVATION AND SUSTAINABLE DEVELOPMENT OF THE PROVINCE OF GALAPAGOS (11 January 2000)

Registro Oficial No. 358, Decree No. 1657

PRELIMINARY SECTION

CHAPTER 1. SCOPE AND PRINCIPLES

Art 3.

. . .

On the basis of the principles of the law, a strategy to achieve the ecological and socio-economic sustainability of human presence in the Galapagos Islands will be prepared. This strategy will serve as a binding referential framework for all plans, projects and activities, that may be developed in, or which are related with, the province. It will be approved by the INGALA Council and will be included in the regional plan, in the terms established in this regulation.

CHAPTER 2. ON PLANNING

Art. 5. The following levels of planning exist in Galapagos:


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a) Provincial or regional planning. The provincial or regional planning will contain the strategy to achieve sustainability and will define the requirements and priorities for the province. Within this framework, the plan will seek the integration and conciliation of the plans of the bodies with responsibilities for planning in their respective areas of jurisdiction within Galapagos, according to the attributions established in the law and other regulatory instruments. The plan will include, among other elements, directives for the protection of the environment and guidelines for achieving the following purposes:

Apply the total control of introduced species, both in protected areas and in urban and rural areas, in accordance with the definition in the glossary of the law;

Improve the levels of education and training of the population;

Promote the social welfare and a lifestyle in accordance with the strategy for sustainability of the region;

Promote the stabilization of population.

b) Planning of government sectors, protected areas and autonomous local authorities. The planning of government sectors, protected areas and autonomous local authorities is that which is carried out by the bodies with specialist responsibilities in terms of the subject matter or the geographical area, as stipulated in the Special Law for Galapagos and other regulations.

Each plan will include an analysis which demonstrates its conformity with the Regional Plan. The INGALA Technical and Planning Committee will evaluate this analysis and, in the event that it considers that there may be inconformity, will provide guidelines for the modification of the plan by the body responsible.

CDF/WWF Biodiversity Vision Annex 3.1 Physical Setting of the Archipelago

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ANNEX 3.1 - Physical Setting of the Archipelago

Note – this information is on a preceding page to avoid disrupting the table.


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Annex 3.1 – Physical Setting of the Archipelago

Island Name1 Latitude

2 Longitude

2 Code

3

Div. Sample

4 Veg.

5

Perm. Isolated

& Aerial6 Area (ha)

7

Coast-line

(km)8

Proximate Isolation

(m)9

Closest Larger Island

10

Dist. from

Center

(km)11

Isla Isabela -0.44506 -91.18803 11 Y Y Y 458812 617.42 74.8Isla Santa Cruz -0.62885 -90.35978 8 Y Y Y 98555 170.51 27600 Isabela 20.6Isla Fernandina -0.38364 -91.52143 12 Y Y Y 64248 114.55 4016 Isabela 112.5Isla Santiago -0.26018 -90.70890 9 Y Y Y 58465 132.18 16860 Isabela 37.2Isla San Cristóbal -0.82014 -89.43485 3 Y Y Y 55808 156.08 66600 Santa Cruz 124.7Isla Floreana -1.29000 -90.43230 5 Y Y Y 17253 61.44 49200 Santa Cruz 84.3Isla Marchena 0.33081 -90.47413 2 Y Y Y 12996 51.39 55800 Santiago 96.6Isla Española -1.37856 -89.68228 6 Y Y Y 6048 42.76 45000 San Cristóbal 132.0Isla Pinta 0.59220 -90.75391 1 Y Y Y 5940 38.95 28800 Marchena 128.1Isla Baltra -0.44781 -90.27278 18 Y Y Y 2619 26.87 361 Santa Cruz 29.3Isla Santa Fé -0.81929 -90.05993 7 Y Y Y 2413 22.84 16653 Santa Cruz 60.1Isla Pinzón -0.61088 -90.66565 4 Y Y Y 1815 16.98 10399 Santa Cruz 18.0Isla Genovesa 0.31732 -89.95957 45 Y Y Y 1410 23.07 46800 Marchena 113.6Isla Rabida -0.41234 -90.70889 10 Y Y Y 499 10.50 4457 Santiago 25.1Isla Seymour Norte -0.39508 -90.28460 28 Y Y Y 184 5.47 1464 Baltra 30.6Isla Wolf 1.37525 -91.81748 47 Y Y Y 134 6.20 140400 Pinta 256.7Isla Tortuga -1.01644 -90.88342 20 Y Y Y 129 9.07 6780 Isabela 66.8Isla Bartolomé -0.28517 -90.55050 46 Y Y Y 124 6.07 310 Santiago 28.2Isla Darwin 1.65656 -92.00537 42 Y Y Y 106 5.41 175200 Pinta 294.3Isla Gardner por Floreana -1.33320 -90.29609 15 Y Y Y 81.2 3.51 7970 Floreana 91.9Islote Cuatro Hermanos Sur -0.86393 -90.77512 39 Y Y Y 72.9 4.40 6154 Isabela 46.0Isla Gardner por Española -1.34335 -89.64356 14 Y Y Y 58.0 3.58 966 Española 132.4Isla Daphne Major -0.42291 -90.37229 19 Y Y Y 33.0 2.14 7600 Santa Cruz 20.8Islote Cuatro Hermanos #2 -0.84844 -90.80045 98 N Y Y 30.4 2.26 3626 Isabela 46.5Isla Eden -0.56010 -90.53763 31 Y Y Y 23.0 2.18 293 Santa Cruz 3.2Isla Caldwell -1.30792 -90.34032 44 Y Y Y 22.8 2.78 2700 Floreana 88.0Isla Sombrero Chino -0.36963 -90.58367 40 Y Y Y 20.8 2.03 112 Santiago 19.9Islote Cuatro Hermanos Oeste -0.84762 -90.81160 97 N Y Y 20.4 1.70 2701 Isabela 47.3Isla Enderby -1.23231 -90.36255 17 Y Y Y 19.3 1.68 2429 Floreana 79.2Roca Bainbridge #3 -0.35055 -90.56583 41 Y Y Y 18.3 1.77 630 Santiago 21.3Islote Venecia -0.51758 -90.47591 27 Y Y Y 13.6 2.27 30 Santa Cruz 5.5Isla Albany -0.17348 -90.84606 43 Y Y Y 12.7 1.55 669 Santiago 54.2Islote Tintorera (Villamil) -0.97201 -90.96107 112 N Y U 12.4 2.64 600 Isabela 68.8Isla Plaza Sur -0.58345 -90.16526 13 Y Y Y 11.9 2.54 302 Santa Cruz 39.9Roca Bainbridge #1 -0.34251 -90.55869 52 Y Y Y 11.4 1.27 1024 Santiago 22.0Islote Campéon -1.23770 -90.38612 16 Y Y Y 9.5 1.28 733 Floreana 79.3Islote Norte de Wolf 1.39235 -91.81792 103 Y Y Y 9 3.06 1000 Wolf 258.3Isla Plaza Norte -0.58047 -90.16176 30 Y Y Y 8.8 2.07 652 Santa Cruz 40.2Roca Beagle Sur -0.41797 -90.63005 78 Y Y Y 8.7 1.30 5264 Santiago 17.9Islote Cráter Beagle #2 -0.28002 -91.35047 11.12 Y Y Y 1. 7 0.71 487 Isabela 96.5Isla Daphne Chica -0.39569 -90.35146 26 Y Y Y 7.9 1.05 10520 Santa Cruz 24.5Isla Sin Nombre -0.67050 -90.58730 65 N Y Y 7.5 0.56 4677 Santa Cruz 16.6Islote Cuatro Hermanos Este -0.84866 -90.75065 99 N Y Y 7.2 1.24 8838 Isabela 43.0Isla Lobos -0.85591 -89.56599 25 N Y Y 6.6 1.50 162 San Cristóbal 111.9Leon Dormido -0.77698 -89.51954 61 N Y Y 5.0 1.06 4608 San Cristóbal 114.4Islote Mosquera -0.40470 -90.27819 38 N Y Y 4.6 1.31 406 Baltra 30.7Isla Caamaño -0.75911 -90.27851 68 Y Y Y 4.5 0.79 1287 Santa Cruz 36.5Roca Bainbridge #6 -0.36760 -90.57029 56 Y Y Y 4.5 0.79 874 Santiago 19.6Roca Beagle Oeste -0.41634 -90.63206 77 Y Y N 4.3 0.70 5102 Santiago 18.2Roca Redonda 0.27478 -91.62304 67 N Y Y 4.3 1.22 24060 Isabela 152.1Roca Bainbridge #5 -0.36409 -90.56643 55 Y Y Y 4.1 0.89 1167 Santiago 19.9Islote Tortuga Oeste -1.02688 -90.88206 115 N U U 3.6 0.82 496 Tortuga 67.6Isla Cowley -0.38466 -90.96236 21 N Y Y 3.5 0.47 3214 Isabela 51.9Islote Guy Fawkes Oeste (largest)

-0.51437 -90.52867 37 Y Y Y 3.4 0.96 2131 Santa Cruz 2.7


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2 Longitude

2 Code

3

Div. Sample

4 Veg.

5

Perm. Isolated

& Aerial6 Area (ha)

7

Coast-line

(km)8

Proximate Isolation

(m)9


10

Dist. from

Center (km)

11

(largest) Roca Bainbridge #4 -0.35869 -90.56446 54 Y Y Y 3.4 0.88 1228 Santiago 20.4Islote Guy Fawkes Sur (2nd largest) -0.51555 -90.52626 36 Y Y Y 3.3 1.01 2343 Santa Cruz 2.5Islote Watson -1.34652 -90.30895 60 N Y Y 3.1 0.82 8012 Floreana 93.0Roca Gordon Este -0.56666 -90.14222 70 N Y Y 2.912 0.45 2990 Santa Cruz 42.2Islote Punta Bowditch Norte -0.53368 -90.51775 32 Y Y Y 2.9 0.92 272 Santa Cruz 0.5Roca Bainbridge #2 -0.34769 -90.55791 53 N Y Y 2.9 0.80 1618 Santiago 21.4Islote Villamil Sureste -0.97571 -90.95623 113 N U U 2.8 1.18 1500 Isabela 68.7Islote Las Bayas Grande -1.22728 -90.44468 86 Y Y Y 2.1 0.97 103 Floreana 77.2Islote Osborn -1.35181 -89.64835 22 Y Y Y 1.7 0.65 430 Española 132.7Islote Cráter Beagle #1 -0.28361 -91.35326 11.11 Y Y Y 1.5 0.62 262 Isabela 96.7Islote Punta Bowditch Sur -0.53600 -90.51754 33 Y Y Y 1.5 0.46 184 Santa Cruz 0.4Islote Guy Fawkes Este -0.49934 -90.51366 35 Y Y Y 1.2 0.55 3448 Santa Cruz 4.3Islote Mao -0.15709 -90.81968 91 Y Y Y 1.2 0.28 200 Santiago 53.7Islote Marielas Sur -0.59876 -91.09086 62 Y Y Y 1.2 0.48 848 Isabela 63.7Islote Fondiadero (Villamil) -0.96705 -90.96321 111 N U U 1.183 0.54 650 Isabela 68.5Leon Dormido Pequeña -0.77862 -89.51896 116 N Y Y 0.975 0.39 4500 San Cristóbal 114.5Islote Cousins -0.23564 -90.57478 69 Y Y Y 0.858 0.32 2000 Santiago 34.0Roca Gordon Oeste -0.56862 -90.14338 72 N U Y 0.828 0.20 2786 Santa Cruz 42.1Roca Dalrymple -0.85580 -89.62718 73 N U Y 0.8 0.03 3933 San Cristóbal 105.4Roca Bainbridge #7 -0.37481 -90.57251 57 Y Y Y 0.796 0.42 1350 Santiago 18.9Roca Este -0.89266 -89.36195 81 N N Y 0.735 0.33 2434 San Cristóbal 134.7Roca Beagle Norte -0.41337 -90.62886 76 N Y Y 0.713 0.57 4805 Santiago 18.2Roca Viuda -0.97528 -90.87177 83 N N Y 0.684 0.23 4389 Isabela 62.4Islote Santa Fé -0.80365 -90.03746 29 Y Y Y 0.671 0.73 17 Santa Fé 61.4Roca Bainbridge #8 -0.37514 -90.57767 58 Y Y Y 0.647 0.40 1148 Santiago 19.1Islote Cráter Cerro Azul -0.91345 -91.38777 11.2 N U U 0.6 1.18 500 Isabela 105.0Islote Muelle (Villamil) -0.96600 -90.95764 114 N U U 0.592 0.62 550 Isabela 68.0Islote Xarifa -1.35729 -89.64427 23 Y Y Y 0.553 0.38 217 Española 133.4Islote Ayora -1.27360 -90.35124 90 Y Y Y 0.5 0.36 600 Floreana 84.0Islote Pitt (nearshore) -0.70109 -89.25226 105 N Y Y 0.5 0.37 41 San Cristóbal 142.2Islote Corona del Diablo Grande -1.21580 -90.42384 48 Y Y Y 0.449 0.38 500 Floreana 76.2Islote Pitt (offshore) -0.70343 -89.24775 66 N Y Y 0.4 0.36 622 San Cristóbal 142.7Islote Oeste -1.34641 -89.66228 24 Y Y Y 0.376 0.17 83 Española 131.1Islote de Canal Sur -0.48495 -90.29345 104 N U Y 0.349 0.27 64 Baltra 25.9Islote Punta Bowditch Este -0.53485 -90.51632 59 Y Y Y 0.346 0.38 113 Santa Cruz 0.6Islote Devine -0.75606 -90.30893 107 N Y Y 0.3 0.38 200 Santa Cruz 33.9Roca Blanca -0.55146 -90.85849 80 N U Y 0.3 0.13 7366 Isabela 37.5Islote Faro (Villamil) -0.96642 -90.96489 110 N U U 0.296 0.17 700 Isabela 68.6Roca Gordon Central -0.56757 -90.14367 71 N U Y 0.259 0.10 2840 Santa Cruz 42.1Islote Marielas Norte -0.59530 -91.09137 63 Y Y Y 0.242 0.21 812 Isabela 63.7La Lobería -0.96358 -90.92635 87 N U U 0.237 0.74 200 Isabela 65.4Islote Guy Fawkes Norte -0.49641 -90.51550 34 Y Y Y 0.235 0.25 3870 Santa Cruz 4.6Islote El Arco 1.64049 -91.99005 108 N Y Y 0.2 1.11 3000 Darwin 291.9Islote Logie -0.25469 -90.57823 92 Y Y Y 0.2 0.21 100 Santiago 32.0Islote Las Bayas Pequeña -1.22798 -90.44284 96 Y Y U 0.145 0.14 94 Floreana 77.3Islote Caleta Tiburón Norte -0.52022 -90.47619 74 N Y Y 0.137 0.10 102 Santa Cruz 5.3Islote Las Cuevas Este -1.26241 -90.36055 101 Y Y Y 0.13 0.20 80 Floreana 82.6Islote Caleta Tiburón Sur -0.52075 -90.47619 75 Y Y Y 0.104 0.10 29 Santa Cruz 5.3Islote Dumb -0.58923 -90.68463 95 N Y Y 0.1 0.27 50 Pinzón 19.1Islote El Torre 1.64905 -92.00684 109 N Y Y 0.1 0.22 200 Darwin 293.7Islote La Ventana 1.36229 -91.82229 89 N Y Y 0.1 1.38 200 Wolf 255.8Islote Las Cuevas Oeste -1.26145 -90.36191 100 Y Y Y 0.1 0.16 50 Floreana 82.4Islote Cráter Beagle #5 -0.28326 -91.34954 11.15 Y Y Y 0.096 0.15 321 Isabela 96.3


133


2 Longitude

2 Code

3

Div. Sample

4 Veg.

5

Perm. Isolated

& Aerial6 Area (ha)

7

Coast-line

(km)8

Proximate Isolation

(m)9


10

Dist. from

Center (km)

11

Islote Corona del Diablo Oeste -1.21668 -90.42350 50 Y Y Y 0.07 0.09 457 Floreana 76.3Islote Noroeste de Santa Fé -0.80522 -90.08704 102 Y Y Y 0.07 0.13 70 Santa Fé 56.7Islote Cráter Beagle #3 -0.27817 -91.35073 11.13 Y Y Y 0.067 0.13 393 Isabela 96.6Islote Marielas Este -0.59479 -91.09077 64 N Y Y 0.067 0.11 738 Isabela 63.6Islote Onan -0.59924 -90.65372 94 N Y Y 0.06 0.31 40 Pinzón 16.3Roca Union -1.03968 -91.09467 82 N N Y 0.05 0.34 2850 Isabela 84.8El Trompo -1.40741 -89.64289 118 N U Y 0.04 0.17 100 Española 137.4Islote Corona del Diablo Este -1.21666 -90.42301 51 Y Y Y 0.04 0.06 100

C. del Diablo G. 76.3

Islote Gran Felipe -0.29115 -90.55425 79 Y Y Y 0.039 0.11 567 Santiago 27.6Roca Rata -0.52245 -90.48578 93 N Y Y 0.035 0.08 40 Santa Cruz 4.2Islote Cráter Beagle #4 -0.27753 -91.35085 11.14 Y Y Y 0.032 0.10 364 Isabela 96.6Islote Caleta Bucanero -0.16806 -90.82739 117 N Y Y 0.02 0.08 70 Santiago 53.3Islote Corona del Diablo Central -1.21613 -90.42350 49 Y Y Y 0.02 0.05 20

C. del Diablo G. 76.2

Islote Espejo 0.30985 -90.40377 88 N U U 0.363* 0.22 Marchena 95.0Islote Este de Darwin -Norte -0.26317 -91.13338 120 N Y Y 0.061* 0.09 20 Isabela 74.5Islote Este de Darwin -Sur -0.26469 -91.13158 119 N Y Y 0.021* 0.05 30 Isabela 74.3Islote La Fé -0.76664 -90.41164 121 N Y Y 1.3* 0.42 3 Santa Cruz 28.3Islote Punta Mangle Norte -0.45191 -91.38605 122 N Y U 0.213* 0.19 60 Fernandina 96.6Islote Punta Mangle Sur -0.45388 -91.38631 123 N Y Y 0.585* 0.31 26 Fernandina 96.6La Botella -1.29039 -90.49910 84 N N U 0.272* 0.19 299 Floreana 83.8Roca Ballena -0.94925 -89.59219 85 N N Y 0.015* 0.05 1087 San Cristóbal 112.9

1 Island names are those listed in Snell et al. (1996), with the additions of Islote La Fé, Islotes Punta Mangle Norte y Sur, Islote Caleta Bucanero, Islotes Este de Darwin Norte y Sur, and El Trompo.

2 Latitude and Longitude values in decimal degrees were calculated by ArcView 3.2 as the centroids of the polygon features for each island. These values differ slightly from those of Snell et al. (1995) because those values were the apparent visual centers of islands. Latitudes and longitudes are correct under the WGS 84 datum. Southern and Western values are negative.

3 Numerical codes used for each island in analyses, databases, and other situations where names are cumbersome. These correspond with the numerical values from Snell et al. (1995). Alpha codes have been replaced with consecutive numeric codes here.

4 “Y” indicates an island with an essentially complete biological diversity survey. Data from those islands are used in later analyses. “N” indicates islands with incomplete or lacking surveys.

5 “Y” indicates an island with at least one species of terrestrial plant, “N” indicates an island without terrestrial vegetation, and “U” indicates an island with undetermined vegetation status. Islands without terrestrial vegetation are included here if they are named in the published literature and/or on official navigational charts.

6 “Y” indicates a landform that is both permanently isolated from others by water at all tides and is always above sea level (not completely awash at high tides). “N” indicates a landform that is either not permanently isolated, and/or is sometimes awash. Landforms that are not permanently isolated or always aerial are included here if they are named in the published literature and/or on official navigational charts.

7 The areas lis ted here are extracted from Snell et al. (1995) for those islands included there. For the additional islands (indicated with an “*”) we calculated areas of their polygons in ArcView 3.2. Polygons representing the islands were digitized from scanned navigational charts (see Snell et al. (1995) for listing) or satellite imagery.

8 Coastline was calculated as the perimeter of the polygons7 using ArcView 3.2.


134

9 Proximate isolation was extracted from Snell et al. (1995) or measured in ArcView 3.2. This measure of isolation is the shortest distance between an island and its nearest larger neighbor. To qualify as a “larger neighbor” and island must be at least twice the size of the other island and not part of the same cluster.

10 This lists the nearest larger neighbor used for measuring proximate isolation.

11 Another measure of isolation is the distance to the geographic center of the archipelago (-0.537°, -90.521). Calculated as the distance from an island’s “center”2 to that of the archipelago in ArcView 3.2.

Figure A3.1: Map of the Galápagos Archipelago.

CDF/WWF Biodiversity Vision Annex 4.1 – IUCN Red Data Book Categories

135

ANNEX 4.1 – IUCN CATEGORIES OF THREATENED SPECIES This annex is a copy of the IUCN criteria for determining categories of Critically Endangered, Endangered and Vulnerable taxa, published in 2000. THE CATEGORIES EXTINCT (EX) A taxon is Extinct when there is no reasonable doubt that the last individual has died. A taxon is presumed Extinct when exhaustive surveys in known and/or expected habitat, at appropriate times (diurnal, seasonal, annual), throughout its historic range have failed to record an individual. Surveys should be over a time frame appropriate to the taxon's life cycle and life form. EXTINCT IN THE WILD (EW) A taxon is Extinct in the Wild when it is known only to survive in cultivation, in captivity or as a naturalised population (or populations) well outside the past range. A taxon is presumed Extinct in the Wild when exhaustive surveys in known and/or expected habitat, at appropriate times (diurnal, seasonal, annual), throughout its historic range have failed to record an individual. Surveys should be over a time frame appropriate to the taxon's life cycle and life form. CRITICALLY ENDANGERED (CR) A taxon is Critically Endangered when the best available evidence indicates that it meets any of the Criteria A to E for Critically Endangered (see below), and it is therefore considered to be facing an extremely high risk of extinction in the wild. ENDANGERED (EN) A taxon is Endangered when the best available evidence indicates that it meets any of the Criteria A to E for Endangered (see below), and it is therefore considered to be facing a very high risk of extinction in the wild. VULNERABLE (VU) A taxon is Vulnerable when the best available evidence indicates that it meets any of the Criteria A to E for Vulnerable (see below), and it is therefore considered to be facing a high risk of extinction in the wild. NEAR THREATENED (NT) A taxon is Near Threatened when it has been evaluated against the criteria but does not qualify for Critically Endangered, Endangered or Vulnerable now, but is close to qualifying for or is likely to qualify for a threatened category in the near future. LEAST CONCERN (LC) A taxon is Least Concern when it has been evaluated against the criteria and does not qualify for Critically Endangered, Endangered, Vulnerable or Near Threatened. Widespread and abundant taxa are included in this category. DATA DEFICIENT (DD) A taxon is Data Deficient when there is inadequate information to make a direct, or indirect, assessment of its risk of extinction based on its distribution and/or population status. A taxon in this category may be well studied, and its biology well known, but appropriate data on abundance and/or distribution are lacking. Data Deficient is therefore not a category of threat. Listing of taxa in this category indicates that more information is required and acknowledges the possibility that future research will show that threatened classification is appropriate. It is important to make positive use of whatever data are available. In many cases great care should be exercised in choosing between DD and a threatened status. If the range of a taxon is suspected to be relatively circumscribed, and a considerable period of time has elapsed since the last record of the taxon, threatened status may well be justified. NOT EVALUATED (NE) A taxon is Not Evaluated when it is has not yet been evaluated against the criteria.


136

THE CRITERIA FOR CRITICALLY ENDANGERED, ENDANGERED AND VULNERABLE CRITICALLY ENDANGERED (CR) A taxon is Critically Endangered when the best available evidence indicates that it meets any of the following criteria (A to E), and it is therefore considered to be facing an extremely high risk of extinction in the wild: A. Reduction in population size based on any of the following: 1. An observed, estimated, inferred or suspected population size reduction of ≥90%

over the last 10 years or three generations, whichever is the longer, where the causes of the reduction are clearly reversible AND understood AND ceased, based on (and specifying) any of the following:

(a) direct observation (b) an index of abundance appropriate for the taxon (c) a decline in area of occupancy, extent of occurrence and/or quality of

habitat (d) actual or potential levels of exploitation (e) the effects of introduced taxa, hybridisation, pathogens, pollutants,

competitors or parasites. 2. An observed, estimated, inferred or suspected population size reduction of ≥80%

over the last 10 years or three generations, whichever is the longer, where the reduction or its causes may not have ceased OR be understood OR be reversible, based on (and specifying) any of (a) to (e) under A1.

3. A population size reduction of ≥80%, projected or suspected to be met within the

next ten years or three generations, whichever is the longer (up to a maximum of 100 years), based on (and specifying) any of (b) to (e) under A1.

4. An observed, estimated, inferred, projected or suspected population size reduction

of ≥80% over any 10 year or three generation period, whichever is longer (up to a maximum of 100 years), where the time period includes both the past and the future, and where the reduction or its causes have not ceased, based on (and specifying) any of (a) to (e) under A1.

B. Geographic range in the form of either B1 (extent of occurrence) OR B2 (area of

occupancy) OR both: 1. Extent of occurrence estimated to be less than 100 km2, and estimates indicating

at least two of a-c: a. Severely fragmented or known to exist at only a single location. b. Continuing decline, observed, inferred or projected, in any of the following: (i) extent of occurrence (ii) area of occupancy (iii) area, extent and/or quality of habitat (iv) number of locations or subpopulations (v) number of mature individuals. c. Extreme fluctuations in any of the following: (i) extent of occurrence (ii) area of occupancy (iii) number of locations or subpopulations (iv) number of mature individuals.

2. Area of occupancy estimated to be less than 10 km2, and estimates indicating at least two of a-c:


137

a. Severely fragmented or known to exist at only a single location. b. Continuing decline, observed, inferred or projected, in any of the following: (i) extent of occurrence (ii) area of occupancy (iii) area, extent and/or quality of habitat (iv) number of locations or subpopulations (v) number of mature individuals. c. Extreme fluctuations in any of the following: (i) extent of occurrence (ii) area of occupancy (iii) number of locations or subpopulations (iv) number of mature individuals. C. Population size estimated to number less than 250 mature individuals and either: 1. An estimated continuing decline of at least 25% within three years or one

generation, whichever is longer, OR 2. A continuing decline, observed, projected, or inferred, in numbers of mature

individuals AND at least one of the following (a-b):

(a) Population structure in the form of one of the following: (i) no subpopulation estimated to contain more than 50 mature individuals,

OR (ii) at least 90% of mature individuals are in one subpopulation. (b) Extreme fluctuations in number of mature individuals. D. Population size estimated to number less than 50 mature individuals. E. Quantitative analysis showing the probability of extinction in the wild is at least 50%

within 10 years or three generations, whichever is the longer (up to a maximum of 100 years).

ENDANGERED (EN) A taxon is Endangered when best available evidence indicates that it meets any of the following criteria (A to E), and it is therefore considered to be facing a very high risk of extinction: A. Reduction in population size based on any of the following: 1. An observed, estimated, inferred or suspected population size reduction of ≥70%

over the last 10 years or three generations, whichever is the longer, where the causes of the reduction are clearly reversible AND understood AND ceased, based on (and specifying) any of the following:






138




of ≥50% over any 10 year or three generation period, whichever is longer (up to a maximum of 100 years), where the time period includes both the past and the future, AND where the reduction or its causes may not have ceased, based on (and specifying) any of the (a) to (e) under A1.


occupancy) OR both: 1. Extent of occurrence estimated to be less than 5000 km2, and estimates indicating

at least two of a-c: a. Severely fragmented or known to exist at no more than five locations. b. Continuing decline, observed, inferred or projected, in any of the following: (i) extent of occurrence (ii) area of occupancy (iii) area, extent and/or quality of habitat (iv) number of locations or subpopulations (v) number of mature individuals. c. Extreme fluctuations in any of the following: (i) extent of occurrence (ii) area of occupancy (iii) number of locations or subpopulations (iv) number of mature individuals.


a. Severely fragmented or known to exist at no more than five locations. b. Continuing decline, observed, inferred or projected, in any of the following: (i) extent of occurrence (ii) area of occupancy (iii) area, extent and/or quality of habitat (iv) number of locations or subpopulations (v) number of mature individuals. c. Extreme fluctuations in any of the following: (i) extent of occurrence (ii) area of occupancy (iii) number of locations or subpopulations (iv) number of mature individuals. C. Population size estimated to number less than 2500 mature individuals and either: 1. An estimated continuing decline of at least 20% within five years or two

generations, whichever is longer, OR 2. A continuing decline, observed, projected, or inferred, in numbers of mature

individuals AND at least one of the following (a-b): (a) Population structure in the form of one of the following:


139

(i) no subpopulation estimated to contain more than 250 mature individuals, OR

(ii) at least 95% of mature individuals are in one subpopulation. (b) Extreme fluctuations in number of mature individuals. D. Population size estimated to number less than 250 mature individuals. E. Quantitative analysis showing the probability of extinction in the wild is at least 20%

within 20 years or five generations, whichever is the longer (up to a maximum of 100 years).

VULNERABLE (VU) A taxon is Vulnerable when best available evidence indicates that it meets any of the following criteria (A to E), and it is therefore considered to be facing a high risk of extinction in the wild: A. Reduction in population size based on any of the following: 1. An observed, estimated, inferred or suspected population size reduction of ≥50%

over the last 10 years or three generations, whichever is the longer, where the causes of the reduction are: clearly reversible AND understood AND ceased, based on (and specifying) any of the following:








of ≥30% over any 10 year or three generation period, whichever is longer (up to a maximum of 100 years), where the time period includes both the past and the future, AND where the reduction or its causes may not have ceased, based on (and specifying) any of the (a) to (e) under A1.


occupancy) OR both: 1. Extent of occurrence estimated to be less than 20,000 km2, and estimates

indicating at least two of a-c: a. Severely fragmented or known to exist at no more than ten locations. b. Continuing decline, observed, inferred or projected, in any of the following: (i) extent of occurrence (ii) area of occupancy (iii) area, extent and/or quality of habitat (iv) number of locations or subpopulations (v) number of mature individuals.


140

c. Extreme fluctuations in any of the following: (i) extent of occurrence (ii) area of occupancy (iii) number of locations or subpopulations (iv) number of mature individuals.


a. Severely fragmented or known to exist at no more than ten locations. b. Continuing decline, observed, inferred or projected, in any of the following: (i) extent of occurrence (ii) area of occupancy (iii) area, extent and/or quality of habitat (iv) number of locations or subpopulations (v) number of mature individuals. c. Extreme fluctuations in any of the following: (i) extent of occurrence (ii) area of occupancy (iii) number of locations or subpopulations (iv) number of mature individuals. C. Population size estimated to number less than 10,000 mature individuals and either: 1. An estimated continuing decline of at least 10% within 10 years or three

generations, whichever is longer, OR 2. A continuing decline, observed, projected, or inferred, in numbers of mature

individuals AND at least one of the following (a-b): (a) Population structure in the form of one of the following: (i) no subpopulation estimated to contain more than 1000 mature

individuals, OR (ii) all mature individuals are in one subpopulation. (b) Extreme fluctuations in number of mature individuals. D. Population very small or restricted in the form of either of the following: 1. Population size estimated to number less than 1000 mature individuals. 2. Population with a very restricted area of occupancy (typically less than 20km2)

or number of locations (typically 5 or less) such that it is prone to the effects of human activities or stochastic events within a very short time period in an uncertain future, and is thus capable of becoming Critically Endangered or even Extinct in a very short time period.

E. Quantitative analysis showing the probability of extinction in the wild is at least 10%

within 100 years.

CDF/WWF Biodiversity Vision Annex 8.1 – Indicators of Ecosystem Function

141

ANNEX 8.1 – PROPOSED GROUP OF INDICATORS OF MARINE ECOSYSTEM COMPOSITION AND FUNCTION The table provides a preliminary list of organisms, which together could serve as indicators of marine composition and function. Note that the “targets of exploitation” column includes species which may be subject to significant indirect effects of exploitation. In particular the workshop group took into consideration not only current exploitation but also the risks of a full-scale sea urchin fisheries, which fishermen have requested, and of open-ocean marine cultivation of algae, which has been mooted and could affect native algae and marine iguanas. The indirectly affected species are in parentheses.

Trophic Structure

Key Stone Important Life history Recruitment

Targets of Exploitation

Indicators of Change Land-Sea Endemic or Rare

Primary producers

Seaweeds Mangroves

Corals Seaweeds

Corals (Seaweeds)

Corals Seaweeds

Seaweeds Mangroves

Seaweeds

Mangroves Grazing invertebrates

Urchins Urchins Urchins Chitons

Urchins Urchins Chitons

Other grazers Iguanas Turtles (Iguanas) Iguanas Iguanas Iguanas Turtles Turtles Piscivores Snapper & Grouper Wahoo Snapper & Grouper Snapper & Grouper Seabirds Snapper & Grouper Seabirds Jacks Seabirds Seabirds Seabirds Wahoo Tuna Wahoo Jacks Planktivores Creole fish Creole fish Black Coral Creole fish Barnacle Blenny Barnacles (Hipponix) Barnacles Black coral Black coral Ascidians (Barnacles) Barnacle Blenny Ascidians Black coral Ascidians Hipponix Benthic predators

Lobsters Conch

Lobsters Conch

Lobsters Conch

Predatory snails Cormorant

Cormorant Conch Cormorant

Snails Whelks Rays

Octopus

Top predators Sharks Marine mammals Octopus Sharks

Sharks Marine mammals

Marine mammals Marine mammals

Marine mammals Detritivores Pentaceraster Sea cucumbers Sand dollars Sand dollars Spider crab

Sea cucumber Spider crab

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• conserving the world’s biological diversity

• ensuring that the use of renewable naturalresources is sustainable

• promoting the reduction of pollution andwasteful consumption

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Logo © 1986, WWF – known internationally as the World Wide Fund forNature, ® Registered Trademark owner.

To conserve the unique biodiversity of Galapagosthrough scientific research and related conserva-tion actions, the Charles Darwin Foundation(CDF), an international, nongovernmental, nonprofit organization, was founded in 1959under the auspices of UNESCO and the WorldConservation Union. The CDF is headquartered inthe Galapagos Islands, where it operates theCharles Darwin Research Station (CDRS) in PuertoAyora on Santa Cruz Island.

The station’s dedicated team of scientists, educators, volunteers, research students andsupport staff from around the world work together to study and protect this precious andunique place.

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Documents

A Biodiversity Vision for the Galapagos Islands