U.S. Dept. of- SHI^ 1 Jun i^^6 ^^4TES O^ Final Environmental Impact Statement for a Possible Regime for Conservation of Antarctic Living Marine Resources June 1978 30°,W 60° W 90° W 120''W 120°E \ ion DEPARTMENT OF STATE FINAL ENVIRONMENTAL IMPACT STATEMENT ON THE NEGOTIATION OF A REGIME FOR CONSERVATION OF ANTARCTIC MARINE LIVING RESOURCES June 1978 3- 3- □ CD r-=l D m /-■»■ Woods i:-.;: ^.„^ano;iraphic insutuiion SUMMARY STATEMENT ■ TYPE : Final Environmental Impact Statement PREPARED BY: Department of State (Attention: Mr. William Mansfield, III Office of Environmental Affairs (Room 7820) Department of State Washington, D. C. 20520) 1. Type of Action: Negotiation of a treaty 2. Brief Description of Proposed Action: The United States will participate in the negotiation of a regime to conserve Antarctic marine living resources. All living resources in waters south of the Antarctic Convergence, except whales and except seals south of 60 degrees south latitude, are affected. The purpose of the regime would be to ensure that any harvesting of Antarctic marine living resources take place in accordance with sound conservation principles. There are no present or proposed commercial fishing operations by the United States in Antarctic waters. 3. Summary of Environmental Impacts and Adverse Environ- mental Effects: Commercial harvesting will result in the continuing and increasing removal of fish and krill from Antarctic waters. Future harvesting may include squid, birds and seals as well. It is anticipated that krill will be the most heavily exploited resource. Populations of organisms which feed on krill and are themselves potentially harvest- able resources will be reduced as krill availability de- clines due to harvesting. The recovery rate of protected baleen whale species may be slowed. Under the proposed conservation regime, harvesting would be regulated to minimize the impact on the Antarctic ecosystem. The adverse impacts of a controlled harvest would be consider- ably less than impacts of uncontrolled harvesting which is anticipated in the absence of an international agree- ment on a conservation regime. Increased harvesting will increase shipping traffic in the Antarctic region. Ship or shore based factory facilities may cause local pollu- tion problems. Increasing human activity will increase disturbance of the Antarctic marine ecosystem which is of 1.1 scientific interest in part because it is the least dis- turbed of marine habitats. The proposed conservation regime, if effective, will moderate adverse environmental impacts in accordance with the objective of maintaining the health and long term productivity of the Antarctic marine ecosystem. 4. Summary of Major Alternatives Considered: Alternatives to the proposed negotiation of a complete conservation regime are: (a) to seek no action to conserve Antarctic living resources, (b) to pursue national action for conservation of Antarctic resources with coordination among nations, (c) to negotiate an international agreement to collect, exchange and analyze data on the Antarctic ecosystem with a commitment to later establishment of a conservation regime, (d) negotiation of a conservation regime, plus imposition of specific conservation measures in the language of the agreement or imposition of a mora- torium on harvesting until such measures are developed, and (e) to negotiate an agreement prohibiting all harvest- ing of living resources in Antarctic waters. 5. Agencies and Parties from Which Comments Have Been Requested; Comments on the draft statement were received from the following: Federal agencies and Offices; U.S. Arms Control and Disarmament Agency Central Intelligence Agency Department of Commerce National Oceanic and Atmospheric Administration Council on Environmental Quality Department of Energy Department of the Interior Marine Mammal Commission National Science Foundation Department of Transportation U.S. Coast Guard XXX Private Organizations; American Cetacean Society, Orange County, Ca. Chapter Center for Law and Social Policy Committee for Humane Legislation, Inc. International Institute for Environment and Development Monitor (Comments endorsed by: Society for Animal Protection Legislation Animal Welfare Institute Humane Society of the United States International Fund for Animal Relief Rare Animal Relief Effort Washington Humane Society International Private Protection League Fund for Animals Defenders of Wildlife Sierra Club Let Live) National Audubon Society New York Zoological Society Individual Comments: David G. Ainley, Ph.D., Point Reyes Bird Observatory J.R. Beddington, University of York, England Hugh H. DeWitt, University of Maine M.D. McWhinnie, Ph.D., DePaul University Donald A. Scott, Oceanographic Services, Inc. Drs. Rudiger Wolf rum and Frederick S. Tipson, University of Virginia 6. Date Draft Statement made available: February 1, 1978 Date Final Statement made available: June 17, 1978 7. Preparation: The draft and final environmental impact statements were prepared by Katherine A. Green, Ph.D. in cooperation with officials of the Department of State, IV TABLE OF CONTENTS Summary Sheet i Table of Contents iv List of Figures vii List of Tables vii List of Appendices viii I. PROPOSED FEDERAL ACTION 1 A. Description 1 B. Concepts Underlying the Proposed Action ... 3 1. Health of the Ecosystem 3 2. Conservation Objectives 4 3. Management Options 6 II. ALTERNATIVES TO THE PROPOSED ACTION 9 A. No Action 9 B. National Action 10 C. Negotiation of an International Agreement for Collection, Exchange and Analysis of Data 11 D. Negotiation of an International Agreement Establishing a Complete Conservation Regime 12 E. Negotiation of an International Agreement Establishing Specific Conservation Measures or a Temporary Moratorium 13 F. A Total Ban on Harvesting 14 G. Area Covered by the Regime 15 V III. RELATIONSHIP OF THE PROPOSED FEDERAL ACTION TO INTERNATIONAL AGREEMENTS WITH IMPLICATIONS FOR THE ANTARCTIC ENVIRONMENT 16 A. Agreements Pertaining to the Antarctic Region 16 B. Agreements Pertaining to the Marine Environment 16 C. Fisheries Bodies 16 IV. RELATIONSHIP OF THE PROPOSED FEDERAL ACTION TO DOMESTIC LEGISLATION WITH IMPLICATIONS FOR THE ANTARCTIC ENVIRONMENT 19 V. DESCRIPTION OF THE ENVIRONMENT 21 A. Physical Environment 21 1. Antarctic Convergence 21 2. Currents 21 3. Ice 22 4. Factors Affecting Primary Production • • 24 B. Species in the Antarctic Marine Ecosystem . 24 1. Krill 25 2. Squid 28 3. Fish 29 4. Birds 29 5. Seals 30 6. Whales 3 0 C. Food Web 31 D. Areas of Special Biological Importance ... 33 E. History of Commercial Harvesting 3 4 1. Whales 34 2. Seals 34 3. Fish 35 4. Krill 35 5. Penguins 3 6 F. Scientific Investigation 36 VI G. Shipping Traffic 37 H. Ecosystem Responses to Disturbance 3 7 I. Potentially Harvestable Resources 3 7 J. No Native Peoples 38 VI. ENVIRONMENTAL IMPACTS OF PROPOSED ACTION AND ALTERNATIVES 3 9 A. Impacts of Commercial Harvesting 3 9 1. Impacts Unaffected by Proposed Action . 3 9 2. Direct Impacts on Exploited Stocks ... 40 3. Indirect Impacts on Dependent Species . 4 2 4. Local Impacts of Harvesting Operations . 44 5. Disturbance of the Marine Ecosystem . .44 B. Environmental Impacts of Proposed Federal Action and Alternatives 45 1. Ban on Harvesting 4 6 2. No Action 4 7 3. National Action 47 4. Data Reporting Now, Conservation Measures Later 58 5. Complete Conservation Regime (Proposed Federal Action) 59 6. Conservation Regime with Specific Conservation Measures or Interim Harvesting Moratorium 6 0 VII. SHORT VS. LONG TERM PRODUCTIVITY 62 VIII. UNAVOIDABLE ADVERSE IMPACTS 63 IX. COMMITMENT OF RESOURCES 6 4 X. CONCLUSIONS 65 XI. CONSULTATION AND COOPERATION WITH OTHERS .... 66 XII. REFERENCES 69 Vll List of Figures 1. Map of the southern ocean 23 2. Food web relationships in the Antarctic marine ecosystem (Green, 1977) 32 List of Tables 1. Common and scientific names of some species occurring south of the Antarctic Convergence. . . 26 2. Level of harvesting operations, feasibility of negotiation, and development and enforcement of conservation measures under the proposed action and alternatives 48 3. Possibility of overexploitation of the various resource populations under the proposed federal action and alternatives 50 4. Changes in abundance of potentially harvestable resources under proposed federal action and alternatives 52 5. Impacts on the marine environment and the health and productivity of the ecosystem under the proposed action and alternatives 54 6. Impacts of proposed federal action and alternatives on collection of scientific data and on the relatively undisturbed ecosystem ... 56 Vlll List of Appendices A - Report of the Ninth Antarctic Treaty Consultative Meeting A-1 B - Transcript of Public Meeting on Antarctic Marine Living Resources, December 20, 1977 B-1 C - Transcript of Public Meeting on Draft Environ- mental Impact Statement for a Possible Conserva- tion Regime for Antarctic Living Marine Resources on February 10, 1978 C-1 D - Interim Report of the Second Special Antarctic Treaty Consultative Meeting and Press Release of the Meeting D-1 E - Role of Krill in the Antarctic Marine Ecosystem . . E-1 F - The Living Resources of the Southern Ocean: Krill F-1 G - The Living Resources of the Southern Ocean: Fish G-1 H - Biological Investigations of Marine Antarctic Systems and Stocks H-1 I - Comments on the Draft Environmental Impact Statement by Federal Agencies, Private Organizations and Individuals I-l J - Department of State Responses to Comments on the Draft Environmental Impact Statement J-1 -1- I. PROPOSED FEDERAL ACTION A. Description The proposed Federal action is the negotiation of a regime to conserve Antarctic marine living resources through conclusion of an international agreement. The international agreement, anticipated to be a treaty, would set forth the objectives of the regime and provide the obligations, func- tions and machinery necessary to fulfill them. The international agreement would provide that: -- the purpose of the regime to be the conservation, including rational utilization, of Antarctic marine living resources ; -- the regime apply to the populations of all living resources of the Antarctic marine ecosystem, defined as all species, including birds, found in marine areas south of the Antarctic Convergence, and their relations with each other and with their physical environment; — the regime not provide for direct regulation of whales and seals already regulated by the International Whal- ing Convention and the Convention for the Conservation of Antarctic Seals; -- any harvesting of Antarctic marine living resources be conducted in accordance with a conservation standard, in- corporating an ecosystem approach, and reflecting the follow- ing elements: a) maintenance of populations or species of Antarctic marine living resources at levels which offer maximum annual produc- tion on a continuing basis; b) maintenance of the balance among the species composing the Antaractic marine ecosystem, taking into account the rela- tionships between harvested and non- harvested species; c) prevention of irreversible changes in the Antarctic marine ecosystem. -2- -- a commission of the Contracting Parties be estab- lished, with an effective decision-making system, to imple- ment the conservation standard and undertake actions neces- sary to ensure the effective operation of the regime, — the commission develop, adopt and revise conserva- tion measures, including total catches in the area as a whole or in subareas defined on the basis of the range of specific populations of Antarctic marine living resources; open and closed seasons and areas, protected species; gear and effort regulation and other steps necessary to fulfill the purposes of the regime; -- a scientific committee be established to provide the commission with independent and expert advice, analyses and recommendations to the commission; — the scientific committee's functions include assess- ment of the status and trends of the populations of Antarctic marine living resources, analysis of data on the effects of harvesting on populations of Antarctic marine living re- sources, assessment of the effects of changes in harvesting levels and proposed conservation measures and identification of research needs and development of proposals to meet such needs; -- the commission publish the reports of the scientific committee; -- a secretariat be established to serve the commis- sion and scientific committee; — contracting parties be obligated to provide data required by the commission and the scientific committee to carry out their functions, including regular and detailed re- porting on harvesting activities carried out by their nationals and vessels; — any harvesting activities be carried out in such fashion to take advantage of opportunities to collect needed data on the impact of harvesting; — contracting parties be obligated to ensure compli- ance with the provisions of the convention and measures adopted by the commission; -- a system of international observation and inspec- tion be established to oversee compliance with provisions of the convention and measures adopted by the commission; -3- — the commission and scientific committee ensure close coordination between their activities and those of the International Whaling Commission and those pursuant to the Convention for the Conservation of Antarctic seals; -- the commission and scientific committee establish cooperative relationships with other international inter- governmental and non-governmental bodies which have responsi- bilities for Antarctic marine living resources, such as FAO, SCAR, and lUCN; -- the budgetary expenses of the commission and other organs of the regime be apportioned equitably among the Con- tracting Parties; — accession to the regime be open to states engaged in research on, or harvesting of, Antarctic marine living re- sources ; -- issues relating to the existence and nature of maritime jurisdiction be treated in such fashion as to permit and encourage participation in the regime by all states en- gaged in research on or in harvesting of Antarctic marine living resources and as to protect the position of the United States regarding non-recognition of claims to territorial sovereignty in Antarctica. B. Concepts Underlying the Proposed Action 1, Health of the Ecosystem One of the purposes of the proposed conservation regime is to ensure that any harvesting of Antarctic marine living resources is conducted in such a fashion as to maintain the health of the Antarctic marine ecosystem. Maintaining the health of the ecosystem implies preserving its overall pro- ductivity over the long term. The structure of the system should be protected by maintaining balance among component populations. The resilience of the ecosystem, or its ability to recover from unusual disturbances such as bad years, should not be reduced. Given the crucial role of krill in the ecosystem, care must be taken that a krill harvest does not cause crashes in the populations of predators which depend on it for food. Of particular concern are baleen whale populations, Increases in those populations depend on reproductive cap- ability and physiological condition of the animals which in turn depend on the abundance of their food supply. -4- All of the breeding stocks for each species of concern should be maintained in a robust condition. In the Antarctic as in other ocean areas, there are natural variations in climate which have negative effects on populations. Unusually warm years, in which pack ice extent is very limited, occur occasionally. These may affect the productivity of plants, and the food supply for zooplankton, fish and birds. In addition some populations may become more vulnerable to predators. Acceptable breeding habitat for seals may be limited. Unusually severe weather, such as prolonged blizzards, may kill all the young of the year for an entire bird rookery. In an undistrubed state, populations can recover from bad years. The size of the breeding stocks of each species should not be reduced to marginal levels where recovery from an unusual year would be slow. For the Antarctic marine ecosystem to remain healthy, its overall productivity should be maintained. The balance between populations and hence the structure of the ecosystem should be maintained both in the system as a whole and in local areas. The inherent resilience of the system, that is its ability to recover from bad years, should not be endangered. 2. Conservation Objectives In addition to maintaining the health of the Antarctic marine ecosystem, other tenets of the proposed federal action include preventing the overexploitation of any Antarctic marine living resource and ensuring that any harvesting of one species does not adversely affect the health of dependent or related species. Understanding of the biology of the Antarctic marine ecosystem is incomplete. Populations of whales are best known , although information is by no means complete. Estimates of abundance, growth and reproduction, and mor- tality rates are available for the harvested species. There is less current information on protected species in the absence of data provided by commercial catches. Abundance estimates are available for seals, but vary over wide ranges. Some information on reproductive rates is also available. However, very little is known of the distribution or survival rates of juveniles, and consequently recruitment into the breeding population cannot be predicted. Since most birds congregate on islands or the coast of the Antarctic continent for breeding, abundance estimates are available for the species and rookeries which have been studied. -5- Fish and squid are known to inhabit the waters, but there is no quantitative information on their abundance or productivity. Even krill, which have been the subject of much scientific and commercial interest, are not well known. Estimates of abundance and of potential yield vary widely. Breeding stocks have not been identified. Some aspects of the physiology of krill such as life span, age of attainment of sexual maturity, and number of reproduc- tive seasons are debated. It is necessary to recognize that information on the Antarctic marine ecosystem is incomplete and to proceed with extreme care in harvesting of any of the resources. Continual monitoring of both the harvested resource and related species is necessary to be able to understand and observe impacts on target stocks and on the rest of the ecosystem. The proposed federal action would require that data from commercial harvesting be both collected and analyzed. It would also encourage scientific investigation. The Antarctic marine ecosystem does have great resource potential and the resources are renewable. Another important objective of the proposed federal action is the maintenance of this potential productivity over a very long term. In managing Antarctic resources there will be trade offs between long term and short term objectives. Inten- sive harvesting to produce high yields for a few years is possible with all the resource stocks, but is inconsistent with a dependable harvest over many decades. There is a real possibility of overexploitation of fish and seal stocks given present technology. As tech- nology for krill harvesting becomes more efficieint, there is also a real possibility of overexploitation of krill even though they are very abundant. In addition, public expectation for krill harvesting is high. The most widely discussed popular estimates of potential krill harvesting are considerably higher than yields which the system could sustain over the long term (Green, 1977) . It is possible that public pressure for a large short term harvest to the detriment of long term yield would be great. One purpose for negotiating a conservation regime for Antarctic living resources is to manage for long term yield of the whole ecosystem. That will entail keeping short -6- term harvesting below the rates which technologically are possible, in order to protect the biology of the system. Since unregulated harvesting would tend to maximize yields in the near future to the detriment of the long term productivity of Antarctic waters, a treaty or regime for long term conservation is needed. 3. Management Options The proposed conservation regime will cover all species of the Antarctic ecosystem except whales which are already covered by the International Whaling Convention and seals south of 60°S latitude which will fall under the Convention for the Conservation of Antarctic Seals now that it is in force. The proposed regime will undertake management of the ecosystem as a whole, i.e. all species together, in cooperation with the bodies governing sealing and whaling. This management approach will be much more effective in maintaining the health of the ecosystem than would be man- agement of the krill population alone. A single species management approach for krill entails a risk of endangering populations of other species which are dependent on krill for food. In addition, it would lack inform.ation on krill losses to predation which will be dependent on the sizes of predator populations. It is possible to undertake an eco- system level approach to resource management in the Antarctic because the ecosystem is relatively simple and the food relationships and interactions among species are understood. Similarly a separate maximum sustainable yield (MSY) determination of potential harvest for each species of commercial interest in Antarctic waters would probably overestimate the total sustainable harvest because of the close interactions among the potentially harvestable resources. Such an approach would not be compatible with maintaining balance among populations or long term pro- ductivity of the ecosystem. Instead the proposed federal action will incorporate an ecosystem management approach in which management goals will include maintaining the long term productivity of the total system and considering balances among the different resource populations. The Antarctic marine ecosystem is defined biologically by species distributions south of the Antarctic Convergence. All waters between the continent and the Convergence are part of the ecosystem. The proposed conservation regime already specifies all waters south of the Convergence as the geographic region to be covered. One alternative would be an agreement on conservation of resources south of 60°S latitude corresponding to the region covered by the Antarctic Treaty and the Convention -7- for the Conservation of Antarctic Seals. However, the Antarctic Convergence falls north of 60°S latitude for most of its extent. There are areas of intense krill con- centrations north of 60°S latitude in the Weddell and Scotia Seas. There are also concentrations of some fish species, elephant and fur seals, and probably some squid species between 60°S latitude and the Convergence. Effec- tive conservation of species not restricted to the pack ice zone must include waters north of 60°S latitude. A second alternative geographic area for a conser- vation regime is choice of a more northern latitude line, such as 50°S latitude, for the northern boundary. That region would include almost all of the waters south of the Convergence. However, it would include subantarctic regions as well and greatly increase the difficulties of negotiating a regime. Although the exact position of the Antarctic Conver- gence does vary, it can be located with simple physical oceanographic measurements. It also has biological mean- ing as a boundary. Of the species which would be included in the proposed management regime only some migratory fish (M. australis and possible some Nototheniids) , fur seals, and probably some squid have distributions which cross the Convergence. For most populations covered by the proposed regime, finding the species would indicate a position south of the Convergence, so that the need to pinpoint its location would be eliminated. Thus the use of the Antarctic Convergence as a boundary for the proposed conservation regime is feasible even though the boundary moves. Within the area to be covered by the regime, it may be possible to manage populations in geographic sectors such as the six whaling regions used by the International Wahling Commission (Chapman, 1974). However, this will not be possible until more information is available on distribution and breeding stocks of the various resource populations. A management approach covering the entire southern ocean, that is all waters south of the Antarctic Convergence, as one region is the best starting point. The proposed conservation regime includes the estab- lishment of a body with the authority to determine conser- vation needs and to regulate harvesting accordingly. Some management techniques which could be used to regulate harvesting include limiting the catch of each species by total catch, by age or size class taken, by geographic area and by season. Allowable gear, total effort or total time spent in harvesting might also be regulated. A concomitant harvest of several species might be contemplated to main- tain balance between populations. Introduction of a har- vestable fish such as salmon into the Antarctic ecosystem -8- to feed on krill and then in turn be harvested has been discussed (Everson, 1977). Such an introduction of species would be subject to regulation by the conservation regime. Requirements to collect and report specific kinds of catch data as well as monitoring of oceanographic para- meters or population sizes, of other than target species might be established. This list of potential management techniques is intended only to provide examples. It is not exhaustive, nor are all the techniques mentioned necessarily to be used. -9- II. ALTERNATIVES TO THE PROPOSED ACTION Negotiation of the proposed conservation regime may- be viewed as one point on a continuum of possible ap- proaches to Antarctic marine living resources. Six gen- eral options can be identified on this continuum. One of these includes the proposed action, the other five repre- sent alternatives to it. They range from a no regime option to seeking a ban on all harvesting, and can be de- scribed as follows : 1. No action to seek to conserve Antarctic marine living resources . 2. National action to conserve Antarctic marine living resources, with coordination among nations taking such action. 3. Negotiation of an international agreement setting forth the obligations and means to collect, exchange and analyze data on Antarctic marine living resources, coupled with a commitment to later establishment of a mechanism to develop and implement necessary conservation measures. 4. Negotiation of an international agreement estab- lishing a complete conservation regime, setting forth the objectives of the regime and providing the obligations, functions and machinery necessary to fulfill them. 5. Negotiation of an international agreement estab- lishing a complete conservation regime and imposing from the beginning specific conservation measures or a moratori- um on harvesting until conservation measures are developed. 6. Negotiation of an international agreement flatly prohibiting all harvesting of Antarctic marine living resources . The proposed federal action under consideration is the fourth alternative. The implications of the six alternatives are discussed below. A. No Action This alternative reflects an assumption that no regu- lation of the harvesting of Antarctic marine living resources is necessary, other than that which takes place or will take place under the International Whaling Convention or the Convention for the Conservation of Antarctic Seals. There is, for instance, no present information that U.S. fishermen -10* plan to fish in Antarctic waters. The presumed potential resources, at least that of krill, are large. Economic factors and operating conditions may serve to limit the extent of commercial harvesting. However, such a laissez-faire approach to harvesting of Antarctic marine living resources would pose the max- imum risks to the Antarctic marine ecosystem and to the long term potential of that ecosystem as a source of sus- tained protein yields. The approach would provide neither for controls on harvesting nor for the means of developing data on the potential resources themselves. The direct dependence on krill for food by most other potential re- source populations makes the dangers of overexploitation particularly acute. A variant of the no action approach would be the post- ponement of efforts to deal with exploitation of Antarctic marine living resources, a no-action-now approach. This option rests on either or both of two premises. One is that the magnitude of the resource is so great that har- vesting need not be regulated for some time to come. The other is that the incentives for an effective system of conservation will be greatest at some point in the future, specifically when large scale harvesting, and its implica- tions, become a reality. There is, however, extensive evidence, based on expe- rience in efforts to conserve marine resources in other areas of the world, that "after-the-fact" regulation is not effective. Sustained unregulated harvesting could have unforeseen, perhaps irreversible impacts on the Antarctic marine ecosystem. Initiation of effective regulation would likely be more difficult once substantial economic stakes in uncontrolled methods of harvesting are created. B. National Action This alternative rests on the assumption that regula- tion by individual countries of vessels and nationals under their jurisdiction engaged in exploitation of Antarctic marine living resources can provide the basis for adequate conservation of the resources. National action could be coordinated through negotia- tion of commitments among the nations involved to regulate harvesting, ranging from a simple undertaking to control harvesting to an obligation to follow specific guidelines in regulating the nationals and vessels under their juris- diction. This alternative could also incorporate informal commitments to share scientific and catch data and to -11- report on harvesting activities with a view to ensuring sufficient coordination to prevent conflicts or overhar- vesting. This alternative would seek to build an essentially voluntary system of conservation in which no nation would be bound internationally. The kinds of obligations this alternative involves could be relatively easy to negotiate. There are, however, a number of practical difficulties inherent in this approach. It would tend to limit effec- tive influence upon the development and implementation of conservation measures to those nations actually engaged in commercial harvesting. The voluntary nature of the system would mean that it would be less stable and could lead to application of inconsistent conservation standards to har- vesting. Its impacts upon the Antarctic marine ecosystem could be quite adverse. C. Negotiation of an International Agreement for Collection, Exchange and Analysis of Data This alternative would concentrate upon the first necessary steps toward providing for adequate conservation of Antarctic marine living resources : establishment of the necessary obligations and means for creating an ade- quate data base. It would involve setting up a scientific and information collecting organization. The alternative is based upon the assumption that it is possible to obtain early agreement on those aspects of a regime which would build the needed data base, with the corollary assumption that negotiation of a full-scale regime could prove suffi- ciently difficult to justify a gradualist approach. This option would satisfy one of the major prereq- uisites for adequate conservation of Antarctic marine living resources, the acquisition of basic scientific data. It might set forth a commitment to establish a system for developing and implementing specific conserva- tion measures at some point in the future. Harvesting would not be regulated until the information gathering system was set up. It could involve a high level of risk to the Antarctic marine ecosystem since there would be no firm guarantees that the necessary standards and machinery to provide for conservation measures would in fact be developed. Delay in addressing the development of conser- vation measures could well make it more difficult to achieve agreement on an effective mechanism. -12- D. Negotiation of an International Agreement Establishing a Complete Conservation Regime This alternative represents the proposed federal action, described in Section I, and rests on the con- clusion that it is urgent to establish an effective system to ensure conservation of Antarctic marine living resources. It also reflects the assumption that the requirements of effective conservation are not incom- patible with properly controlled harvesting of Antarctic marine living resources. In addition, under this approach, harvesting could take place in the interim period after the conclusion of the regime and prior to the development of conservation measures pursuant to the regime. The question of the treatment of harvesting in this interim period can be dealt with in two ways, which com- prise variants or sub-options of the proposed federal action. The first sub-option would be not to address specifically the interim period. This variant assumes that technological and other limitations will ensure low harvesting levels for the next several years. It rests on the view that a comparison of projected initial levels of harvesting with projected potential of stocks of commercial interest indicates that specific limita- tions in the interim period are not required. In other words, this sub-option assumes that harvesting will not reach levels which would pose threats to target species, to dependent species or to the ecosystein as a whole prior to the development of the regime's full capability to identify conservation needs and apply conservation measures. In the event that development of the conser- vation machinery requires many more years than presently anticipated, this variant would involve some risk of overexploitation . The second sub-option involves negotiation, as a supplement to the international agreement, of interim measures to apply during the period after conclusion of the regime and prior to its becoming operational. Interim measures would include mechanisms for beginning to build the data base necessary for effective operation of the regime and possible interim ceilings on harvesting, at least for krill. These interim measures would be designed to reflect a conservative approach to harvesting activities during the period prior to the full operation of the system set forth in the conservation regime. -13- The first of these sub-options would likely provide the necessary margin of safety in avoiding harmful impacts upon the Antarctic marine ecosystem, and would be the easier to negotiate. The second sub-option would be more difficult to negotiate but would reflect a more cautious and conservative approach to exploitation of resources, such as krill, with which there is little international experience. For this reason, the second sub-option or variant, providing for the interim measures approach described above, is preferred. E. Negotiation of an International Agreement Establishing Specific Conservation Measures or a Temporary Moratorium This alternative would involve negotiation in the international agreement of specific conservation measures for some or all Antarctic marine living resources or a moratorium on harvesting until such measures can be devel- oped. It would imply a conservation regime similar to that in the proposed federal action. The alternative would differ from the proposed federal action in one of two ways : -- the agreement would set forth detailed conser- vation measures for some or all species, rather than provide for interim catch limitations or for other specific initial conservation measures to be developed by the organization established pursuant to the regime, or — the agreement would impose an obligation to abstain from harvesting until the conservation regime became fully operative rather than permit harvesting to take place while conservation measures were being developed. This approach would pose fewer risks to the Antarctic marine ecosystem than the previous alter- natives, including the proposed federal action. However, it suffers from several major drawbacks. There is insufficient data upon which to base specific conser- vation measures. An attempt to do so ab initio could significantly decrease incentives for nations with harvesting interests to participate in the conservation regime. A moratorium on harvesting until sufficient -14- data is developed suffers to an even greater degree from the same negotiating disadvantages. Since a small controlled catch of krill and other species would provide data on size and distribution of stocks which are required for proper management but not presently available, a moratorium would increase the difficulty of determining necessary conservation measures. It would be extremely difficult to determine when sufficient information existed to justify lifting the moratorium. A fixed time period for the moratorium might be considered equally arbitrary. An effective conservation regime should include all states engaged in harvesting. This alternative risks losing the participation of such countries, thus leading to a less effective system than that of the proposed federal action. F. A Total Ban on Harvesting Maintenance of the Antarctic ecosystem in its present state would require a prohibition on all har- vesting of Antarctic marine living resources. From the conceptual point of view, this approach obviously offers the maximum degree of protection to that ecosystem. Seeking a ban on harvesting would ultimately represent a judgement that the health of the Antarctic marine ecosystem can be ensured only by its total insulation from human activity. By contrast, the proposed federal action is based upon the assumption that properly controlled harvesting of Antarctic marine living resources is consistent with maintenance of the health of the Antarctic marine ecosystem. Further, a total ban on harvesting stands no chance of being negotiated. States interested in commercial harvesting would have no incentive to accept this alternative, and as a practical matter, it would be equivalent to no regulation whatsoever. Preservation of the Antarctic marine ecosystem as a sanctuary or a relatively undisturbed natural habitat free from commercial harvesting is an unobtain- able objective. -15- G. Area Covered by the Regime There is a subsidiary set of alternatives concern- ing the geographic area to be covered by the regime. The conservation regime of the proposed federal action would apply to the species which comprise the Antarctic marine ecosystem. The Antarctic marine ecosystem is generally considered to be geographically defined by the Antarctic Convergence which shifts in space on a seasonal basis. Linking the regime to the species which are found south of the Convergence best provides for coverage of the full ecosystem. There are two alternatives to this approach based on geographic limits, which would provide for enforce- ment of conservation measures. The first is to limit the regime to the area of the Antarctic Treaty, south of 60°S latitude. Such a result could be the easiest to negotiate, at least among the Antarctic Treaty Parties. However, it would seriously compromise the effectiveness of conservation measures. Since the Antarctic Convergence lies significantly north of the Treaty area in a number of places, areas of important concentrations of Antarctic marine living resources would not be covered. A second possibility would be to fix a geographic limit for the applicability of the regime sufficiently far north to ensure that all areas south of the Convergence were covered. The difficulty with this approach is that it would cover species not part of the Antarctic marine ecosystem and thus create difficult technical and political obstacles to negotiating the regime . -16- III. RELATIONSHIP OF THE PROPOSED FEDERAL ACTION TO INTERNATIONAL AGREEMENTS WITH IMPLICATIONS FOR THE ANTARCTIC ENVIRONMENT A. Agreements Pertaining to the Antarctic Region The Antarctic Treaty reserved for peaceful uses the Antarctic continent and ice shelves, south of 60°S latitude. It encourages scientific investigation and provides for exchange of information. It further provides nothing in the treaty shall prejudice or in any way affect the rights, or the exercise of rights of any State under international law with regard to the high seas within the Treaty area. In addition to the Antarctic Treaty itself, the United States is a party to several agreements which directly con- cern the environment of Antarctica. The Agreed Measures for the Conservation of Antarctic Fauna and Flora (not yet in force, but effectively adhered to as voluntary guidelines by the Treaty Parties) provide a stringent system of protection of native mammals, birds and plants and establish certain "Specially Protected Areas" which are to be accorded special protection in order to preserve their unique natural ecological characteristics. The Agreed Measures pertain to areas and species on the Antarctic continent and pack ice. The International Whaling Conven- tion regulates whaling in Antarctic waters. The Convention for the Conservation of Antarctic Seals, opened for signa- ture in 1972, entered into force on March 11, 197 8. It applies south of 60° latitude. The Convention on International Trade in Endangered Species of Wild Fauna and Flora entered into force 2 June 1975. The Convention provides for the identification of endangered species and the prohibition in international trade in such species or products made therefrom. It applies to two species of whale — the blue and humpback whale -- found in Antarctic waters. These whales are also protected species (no commercial harvesting) under the International Whaling Convention. B. Agreements Pertaining to the Marine Environment In addition to the foregoing international agreements , a number of international conventions dealing with protec- tion of marine environment, primarily from vessel-source pollution, would apply to Antarctic waters. Regulation of discharges from vessels and provision of liability for damages and clean-up costs for spillages of oil into the -17- the marine environment are the subjects of the following conventions negotiated within the Inter-governmental Maritime Consultative Organization (IMCO) : — The International Convention for the Prevention of Pollution of the Sea by Oil, 1954 as amended in 1962 and 1969. (Ratified by the U.S., entered into force 20 January 1978) . — The International Convention for the Prevention of Pollution from Ships, 1973. (Not ratified by the U.S., not yet in force) . — The International Convention on Civil Liability for Oil Pollution Damage, 1969. (Not ratified by the U.S., entered into force 19 June 197 5) . — Protocol to the International Convention on Civil Liability for Oil Pollution Damage, 1979. (Not ratified by the U.S., not yet in force). — International Convention Relating to Civil Liability in the Field of Maritime Carriage of Nuclear Material, 1971. (Not ratified by the U.S., entered into force 15 July 1975). — The International Convention on the Establishment of an International Fund for Compensation for Oil Pollution Damage, 1971. (Not ratified by the U.S., not yet in force). — Protocol to the International Convention on the Establishment of an International Fund for Compensation for Oil Pollution Damage, 1971. (Not ratified by the U.S., not yet in force) . International regulations of the intentional disposal of matter at sea (which again applies to Antarctic waters) is provided for in the Ocean Dumping Convention (the Conven- tion on the Prevention of Marine Pollution by Dumping of Wastes and Other Matter, of 1972) which the U.S. has ratified and which entered into force on August 30, 197 5. The commitments which would be undertaken by parties to the regime proposed in the federal action would be entirely consistent with the obligations under the international agreements listed above. The legal instrument setting forth activities pursuant to the regime with those pursuant to the International Whaling Convention and the Convention for the Conservation of Antarctic Seals (when in force) . In addition, it would be desirable to provide specific refer- ence to the Agreed Measures for the Conservation of Antarctic -18- Flora and Fauna in the legal instrument to ensure that measures pursuant to the Agreed Measures are properly taken into account. C. Fisheries Bodies There are several fisheries bodies with which a regime to conserve Antarctic marine living resources might have specific relationships. These include the Regional Fisheries Advisory Commission for the Southwest Atlantic, the Indian Ocean Fishery Commission, the Indo-Pacific Fisheries Council, the International Commission for the Southeast Atlantic Fisheries, and the Permanent Commission of the Conference on the Use and Conservation of the Marine Resources of the South Pacific. The first three bodies were created under the auspices of the United Nations Food and Agriculture Organization (FAO) , the other two purusant to separate international agreements. The proposed federal action would provide for estab- lishment of cooperative relationships, where appropriate, with those bodies. This could be accomplished through the close working relationship anticipated with FAO, supplemented if and where necessary with ties to any of the individual fisheries bodies. -19- IV. RELATIONSHIP OF THE PROPOSED FEDERAL ACTION TO DOMESTIC LEGISLATION WITH IMPLICATIONS FOR THE ANTARCTIC ENVIRONMENT There are two domestic acts which bear upon species covered by the conservation regime set forth in the pro- posed Federal action: the Endangered Species Act of 1973 and the Marine Mammal Protection Act of 1972. The Endangered Species Act of 1973 incorporates and expands authority under previous endangered species legis- lation to identify fish and wildlife threatened with extinction and to prohibit their importation into the coastal States. The 1973 legislation gives consideration to all animal life, not only the vertebrates, mullusks and crustaceans included under the previous law, and recognizes the importance of wild plants as well as animal species. It provides for protection of "threatened" as well as "endangered" species, permitting preventive action before a critical stage is reached and thereby enhancing the like- lihood of successful recovery. It authorizes a grant program to assist state endangered species programs, and provides for Federal protection of resident species where states are unable to do so. It requires coordination among all Federal agencies whose activities may impact threatened or endangered species or their habitats, and directs these agencies to use their other authorities in further- ance of the purposes of the Act. Species found in Antarctic waters listed as endangered under the U.S. Endangered Species Act are the blue whale, humpback whale, sei whale and sperm whale. (The sperm whale is not listed as endangered in the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) . Sei whales other than the North Pacific stock, are not listed as endangered under CITES, but are listed as stocks to which special controls should apply.) The Marine Mammal Protection Act of 1972 gives special domestic protection to marine mammals, including whales. The Act prohibits the taking or importation of such mammals and their products by U.S. citizens. The Secretary of Commerce or Interior, depending on the species, can waive this prohibition only if he receives scientific evidence that the waiver and the regulations on takings that he must develop after a formal hearing, will not be to the disadvantage of the species or stock to be taken. The Act calls for negotiations to develop international -20- programs for the protection and conservation of marine mammals covered by the Act. The conservation regime set forth in the proposed Federal action would be consistent with the purposes of both the Endangered Species Act and the Marine Mammal Protection Act. -21- V. DESCRIPTION OF ENVIRONMENT A. Physical Environment Surrounding the Antarctic continent is a ring of ocean contiguous with the Atlantic, Pacific and Indian Oceans to the north. The Antarctic Convergence is the northern boundary of Antarctic waters. Waters north of the Convergence are subantarctic . The average ocean area south of the Convergence is 38 x 10^ km^ (El-Sayed, in press) . 1. Antarctic Convergence The Antarctic Convergence is a boundary between different water masses. South of the Convergence water surface temperature drops 2 to 3°C. The exact geographic location of the Convergence varies with changing thermo- cline and wind stress conditions, but it can be located with temperature and salinity measurements. The Antarctic Convergence also known as the Antarctic Polar Front Zone, is not a very sharp boundary, but actually is a transitional zone between water masses. It is a charac- teristic of surface water masses, having a depth of up to 600 meters. The deep northward flowing Antarctic bottom water flows underneath the Convergence (Gordon, 1971) . The Antarctic Convergence is not a boundary for deep living organisms such as bathypelagic fishes. A map of the southern ocean showing the position of the Convergence and other features is given in Figure 1. The Convergence lies approximately along 50 S latitude in the Indian and Atlantic Ocean sectors. In the Drake Passage it lies approximately halfway between the tip of the Antarctic peninsula and the tip of South America. In the Pacific sector it lies approximately between 55°S and 60°S latitude (Mackintosh, 1972) . The Convergence is a physical oceano- graphic boundary. It is also a faunal boundary. Biological communities south of the Convergence are distinct from those north of it (Marr, 1962; Hasle, 1968). 2. Currents Surface currents flow latitudinally in the southern ocean. Near the continent water flows toward the west and is known as the East Wind Drift. North of approximately 60OS latitude water flows toward the east and is known as the West Wind Drift or the Circumpolar Current. There is a -22- gyre circulation pattern in the Weddell Sea extending from the Antarctic peninsula to almost 30°E longitude (Foster, in press) which seems to coincide with some species distri- butions (Everson, 1977) . There are other gyres in the Bellingshausen and Ross Seas. .Plankton are transported by water currents. 3. Ice Seasonal variation in the ice cover characterizes the southern ocean. Large icebergs form from calving of the tongues of glaciers. Since bergs extend deep into the water, their movements are determined by patterns of water currents. The much more extensive sea ice which covers enormous areas of ocean surface is formed by the freezing of sea water and is known as pack ice. It averages about 1 meter in thickness, and its movements are determined by wind patterns. The pack ice extends northward from the continent with a maximum areal extent of approximately 25.5 x 10" km'^ in the winter and a minimum of about half that or 13 x 10 km in the summer (Mackintosh and Brown, 1956). Thus the Antarctic continent appears to pulsate. The total extent of the pack ice varies from year to year. The outer edge of the pack ice is always several degrees of latitude inside the Antarctic Convergence (Mackintosh, 1972). The pack ice provides habitat for seals and penguins. Most of the pack ice melts during the summer and so has a maximum age of one year. Some multi-year ice is found in the Weddell Sea where gyre circulation prevents it from moving northward. Areas of pack ice in February and March (late summer in the southern hemisphere) in the eastern Ross Sea and along much of the coast of the Antarctic continent presum- ably are population centers for pelagic seals (Gilbert and Erickson, 1977) . As the pack ice expands and contracts, it covers and un- covers krill and other nonmigrating populations. There is intense biological activity at the edge of the pack ice where primary productivity is high. Herbivores are active and feeding on plankton is intense (Andriashev, 1968). Whale migration routes are related to pack ice distri- bution. In the summer whales are found in open water, with the most southern portions of the stocks approaching the edge of the pack ice. South Atlantic Ocea Antarctic Con }iergence Falkland Islandi /B^)ingshiusen b; Sed, \\ Antarctica -^^ The Southern Ocean k TT A \ \ Oes de Ke'goe'en ti-. \ McDonald Islands . \ Heard Isfand y J Indian Ocean ■A- Ice Limit ^Wacoua'ie Island South Pacific Ocean CampbelUsland ' Auckland Islands 'Aniifyides Islands Boundary represenladon is nol necessarily aiithoMaliwe Bounty Islands ' 180 So New /Zealand Australia -24- 4. Factors Affecting Primary Production Antarctic waters are rich in nutrients. Circumpolar upwelling introduces nutrients into the euphotic zone throughout the year (Gordon, 1971) . Water temperature is uniformly cold. The available light varies seasonally with the polar light regime and is closely correlated with primary production (El-Sayed and Mandelli, 1965). Primary production is concentrated in the summer season, along the ice edge and in coastal regions (Balech, et al, 1968; El-Sayed, 1970) . Because the early scientific investigations were concentrated in the most productive areas, a misleading picture of very high productivity for Antarctic waters was developed. Estimates have since been revised downward. On a yearly average basis over all southern ocean waters, in- cluding the less productive open ocean regions, total pro- ductivity is similar to that in other oceans (Holm-Hansen et al, 1977) . Food chains are shorter in the Antarctic than in other oceanic regions. Plants are consumed by krill (the dominant herbivores), which are in turn eaten by fish, birds, and mammals. But because food chains are shorter, a greater proportion of the productivity reaches the vertebrate trophic levels, and there is an unusually high density of birds and mammals in Antarctic waters. Variation in sea ice cover affects light available to the water column because ice screens out light. Primary produc- tion in the winter is extremely low. B. Species in the Antarctic Marine Ecosystem There is a distinct fauna south of the Antarctic Con- vergence. Phytoplankton consist primarily of diatoms south of the Convergence, but include a great many dinof lagellates north of it. Zooplankton species are different north and south of the Antarctic Convergence. Euphausia superba and E. crystallorophias are only found south of the Convergence, E. superba in oceanic regions and E_^ crystallorophias near the continent (Marr, 1962) . The term krill refers to the group of euphausiid crusta- ceans which are the food of large baleen whales. Species of krill are listed in Table 1. By far the most abundant of these species is E_^ superba. -25- Krill are omnivores, eating other zooplankton as well as phytoplankton. Krill are usually considered to be about half of the zooplankton population (Gulland, 1970) . Other zooplankton include copepods, araphipods , and chaetognaths (Hopkins, 1971) . Krill play a crucial role in the Antarctic marine food web because of their great abundance and because they are the main prey item for most of the predators in the ecosystem. Squid are probably abundant in the Antarctic (Appendix F. BIOMASS, 1977). Squid eat krill and fish. Many fish eat krill. Birds, predominatly Adelie penguins, also eat krill (Prevost, in press) . Other Antarctic birds include other penguins, petrels, albatrosses, and skuas. Crabeater seals, a krill eating species, are the most abundant of Antarctic seals. Leopard, Ross, Weddell, elephant and fur seals are also present. The baleen whales which migrate into the Antarctic, fin, blue, sei, humpback, and minke whales, feed extensively on krill. Southern right whales are occasionally found south of the Convergence. The toothed whales, sperm and killer whales and some small ceta- ceans, are probably resident throughout the year. Common and scientific names for species discussed in describing the Antarctic marine ecosystem are listed in Table 1. At the present state of biological knowledge, standing stocks of resource populations other than whales in Antarctic waters are not well known. Abundance estimates for seals and penguins still vary considerably among investigators. The overall abundance of fish and cephalopods is virtually unknown. Abundance estimates and educated guesses on stock sizes for potential resource populations are discussed in Appendices E, F, G and H. 1. Krill Krill have a circumpolar distribution, but are most abun- dant in the Weddell Sea, the East Wind Drift, the Weddell Drift and the South Georgia region (Nemoto, 1968; Marr, 1962; Mackintosh, 1973) . Krill are found both under ice and in open waters. E. superba collect in large amorphous swarms varying from a few to several hundred meters across. Swarms are ap- parently composed of individuals of a single age class. Swarms occur at varying depths from the water surface down to about 100 m. Not all krill are found in swarms. Even in regions of greatest abundance krill distribution is patchy -26- Table 1. Common and scientific names of some species occurring south of the Antarctic Convergence. Zooplankton Krill Euphausia superba E. cry stal lorophias E . tria can tha E . fri gi da Thysanoessa macrura T . vicina Fish Antarctic cod Notothenia rossii N. gibber i frons N . cor i i ceps N . magel lanica N . negl ecta Antarctic tooth fish Dissost ichus mawsoni Patagonian tooth fish D. eleginoides southern blue whiting Micromesistius australi s or southern poutassou Antarctic silverfish Pleuragramma anta ret icum icefish Channichthys rhinoceratus Chaenocepha lus aceratus Champsocephalus gunnari Pseudochaenichthy s georgianus -27- Table 1 (Continued) Squid giant squid other squid Birds Adelie penguin Seals crabeater seal leopard seal Ross seal Weddell seal elephant seal fur seal Whales fin whale blue whale sei whale minke whale humpback whale southern right whale sperm whale killer whale Mesonychoteuthi s hami Itoni Moroteuthi s ingens Conatus fabricii Pareledone sp . Pygoscel is adeliae Lohodon card nophagus Hydrurga leptonyx Omatophoca rossi Leptonychotes wedde 1 1 i Mirounga leonina Arctocephalus gazella Ba 1 aenoptera phy sa 1 us B . musculus B . boreal i s B. acutorostrata Megaptera novaeangl iae Euba laena aus tra 1 i s Physeter catodon Orel nus orca -28- (Appendix D, Everson, 1977; Tetra Tech, 1978) . Because of its swarming habit, E_^ superba is the predominant krill species in the diets of mammals and penguins, and is the only krill species of commercial importance. Estimates for krill standing stock vary almost tenfold, from about 180 to 1350 million metric tons (Appendix E, Green, 1977) . The circumpolar krill population may be a single breed- ing stock or may be composed of several separate stocks (Mackintosh, 1973; Makarov, 1974). More recent studies indi- cate that the breeding area has spread geographically since spawning females have been taken from the Scotia Sea, Belling- shausen Sea and Bransfield Straits. Spawning occurs at the surface from late November through late March. Early larvae are found in the East Wind Drift, the Bransfield Strait, the Scotia Sea, and the Weddell Drift (Everson, 1977; Tetra Tech, 1978). The Discovery expeditions collected very few eggs, and those only in the Scotia Sea (Marr, 1962). Krill are long lived in comparison with zooplankton in warmer waters. However, there is still considerable doubt about total life span and age at which sexual maturity is attained. Krill may not breed until the beginning of their third or fourth summer and may breed for one or two seasons (Eraser, 1937; Bargmann, 1945; Ivanov, 1970; Everson, 1977) . Even though there is considerable scientific and commer- cial interest in krill, knowledge of physiology, breeding stocks, replacement rate and abundance is still incomplete. 2. Squid It is possible that squid in the southern ocean are con- centrated near the Antarctic Convergence (Appendix Hf BIOMASS, 1977). However, squid serve as food for sperm whales, seals and fish whose distribution extends south toward the continent. No direct abundance estimates for squid or other cepha- lopods are available for Antarctic waters, mainly due to inadequate sampling techniques (Everson, 1977) . There are squid fisheries in the waters near New Zealand and South America, but none in Antarctic waters. The "giant squid" Mesonychoteuthis hamiltoni may be of commercial potential (Everson, 1977) . Squid are major predators on krill. Larger individuals feed on fish or other squid. -29- The squid group, like the fish group, are krill predators and themselves serve as food for other predatory species as well as potentially harvestable resources. The presence of potentially harvestable populations are several trophic levels i.e., herbivores and primary and secondary carnivores, is a characteristic of the Antarctic ecosystem. 3. Fish I Fish stocks are thought to be low in the Antarctic (Gulland, 1970) . In contrast to other oceans, there are not dense shoals of obligate pelagic fish in Antarctic waters. Pelagic fish present in the waters are Myctophidae, the Noto- theniid genera Pleuragramma and Dissostichus, and the Gadoid genus Micromesistius . Although the Myctophidae may be abun- dant in the open ocean, they apparently do not form dense enough concentrations to support a fishery (Appendix H, Everson, 1977) . The Antarctic cod or Nototheniids are distributed in coastal areas around islands such as the Scotia Arc, Ker- guelens, Crozet, Bouvet, and in the Magellanic and New Zealand regions. The latter may migrate south to the Ross and Scotia Seas in the summer, but most of the fish populations are probably resident in Antarctic waters. The southern blue whiting Micromesistius australis from the Magellanic and New Zealand regions does migrate into the Antarctic zone only during the summer to feed. The Antarctic tooth fish and Antarctic silverfish are found around the continent over continental shelves. Antarctic cod and some ice fish are being harvested commercially. Other fish species may consti- tute potentially harvestable resources. 4. Birds The Antarctic Convergence is not a barrier for birds because of their mobility. However, breeding of penguins, albatrosses and petrels, cormorants, skuas, ducks, gulls and terns occurs along the coast of the Antarctic continent or on islands between 50°S and 65°S latitude. South Georgia and Kerguelen have the greatest diversity of breeding birds (Watson, et al, 1971; Everson, 1977) . Migratory birds are present in the Antarctic ecosystem, either south of the Convergence or on subantarctic islands, for only part of the year for feeding and breeding. The eco- system provides habitat for such bird populations. When they move northward, they export some material from the Antarctic ecosystem. -30- Birds eat krill, squid, and fish. Adelie penguins are the most abundant species (measured by biomass) south of the Convergence. They consume very large amounts of krill in proportion to their body weight (Prevost and Sapin- Jaloustre, 1965; Prevost, in press) . They are eaten by leopard seals and killer whales (Gilbert and Erickson, 1977) . 5. Seals The crabeater seals, whose population may be as high as 30 million individuals, are the most abundant seal in the Antarctic region and the most abundant seal in the world. They are residents of the pack ice region and feed mainly on krill. Other pack ice seals are the leopard and Ross seals. Pack ice seals breed in the pack ice region, usually south of 60°S latitude. Distribution of juveniles of these popula- tions is unknown and they may occasionally range north of 60°S. Weddell seals inhabit coastal areas of fast ice. Ross and Weddell seals eat fish and squid as well as some krill. Leopard seals are predatory, eating mainly krill but also some fish, the young of other seal populations and occasionally penguins . Elephant seals and fur seals breed on islands near the Convergence. The main population of the fur seal Arctocephalus gazella is at South Georgia. Other fur seals breed north of the Convergence. Elephant seals eat fish and squid; fur seals eat mainly krill. Elephant seals and fur seals are also resident in the Antarctic region (Erickson, et a_l, 1971; Erickson and Hofman, 1974; Appendix H, BIOMASS, 1977). 6. Whales Whales are circumpolar in distribution. The baleen whales, or fin, blue, sei, humpback and minke whales, migrate into Antarctic waters during the austral summer to feed on krill. Southern right whales are occasionally found south of the Convergence. Baleen whales are found south of the Convergence from November through March. Individual animals average about four months feeding in the area. Whales feed in open water. The larger species and larger individuals within species penetrate further south, approaching the edge of the pack ice. While feeding intensively in Antarctic waters, baleen whales increase about 50% in weight during the sumirier season (Mackintosh and Brown, 19 74; Laws, 19 77a; Appendix H, BIOMASS, 1977) . Sperm whales, killer whales and some small cetaceans are probably resident in the Antarctic region. A third of the exploitable male sperm whale stock is assumed to feed south -31- of the Convergence. Toothed whales eat fish and squid, possibly penguins and juvenile seals (Mackintosh and Brown, 1974; Laws, 1977a; Appendix H, BIOMASS, 1977). Fin, blue, humpback and southern right whales are pro- tected from harvest. The present population size of fin whales is about 20% of the original stock; blue whales are about 5%, and humpback whales about 3%. The sperm whale population is now about half of initial stock size and is now harvested. Sei whales which are harvested in some areas are presently slightly above half of initial popula- tion size. Minke whales, also harvested, are still as abundant as they were a century ago (Laws, 1977a). As population size of baleen whales species has de- creased, there have been changes in other population para- meters. Sexual maturity is reached at a younger age now than when stocks were larger. The pregnancy rate is also increasing. These changes combine to increase the growth rate of the total population at low stock levels (Everson, 1977) . A growing population requires more food than a stable population of the same size. Thus it is possible that if whales were to expand populations to former levels, at some point their food requirements would be greater than the total food requirement of the original stock. C. Food Web The importance of krill in the Antarctic marine ecosystem cannot be overemphasized. They are the most abundant zoo- plankton species. They are the main herbivore in the Antarctic marine ecosystem. Krill are the main food item for fish, squid, penguins, crabeater and leopard seals, and fin, blue, sei, humpback, and minke whales. Krill are present in the diet of other seals as well. Killer whales and sperm whales, in consuming fish and cephalopods and some seals, are only one step removed from krill in the food chain. A food web diagram for the Southern Ocean marine ecosystem is presented in Figure 2. The dependence of many predators on one prey group, krill, is a very unusual ecological situation. The swarming habit of krill allows even very large predators to feed on them efficiently. Removal of krill from the ecosystem by commer- cial harvesting will compete directly with those predator populations feeding in the same geographic area and indirectly with other predators as overall abundance changes. -32- 4-1 m >^ w O u (U •H CJ u CO •u C < 4-1 a CO c o 0) >^ o o CNI 0) •H -33- If krill should be so depleted that some other group, such as copepods , became the most abundant zooplankton, a difference in overall food web structure for the ecosystem could result. Since copepod populations are dispersed, they would not support the same kind of large predators which krill can support by swarming. A hypothetical copepod- myctophid food chain might contain the same amount of carbon as the krill food web does now, but much more dispersed, so that the resources would not be as readily available, either to large predators or to commercial harvesting, as they are in the present system. Crabeater seals are the biggest consumer of krill, taking about 106 million metric tons annually. Squid may take 100 million tons and fish 60 million tons although those esti- mates are not reliable, since fish and squid abundance esti- mates are speculative. Baleen whales consume about 43 million tons annually at present, birds about 14 million tons, and other seals about 3 million tons. Estimated annual total predation on krill is about 340 million tons. Krill harvested will come from the total now consumed by predators, not from some hypothetical part of the stock which is not utilized by predators (Appendix E, Green, 1977) . At initial stock levels, baleen whales consumed about 150 million tons more krill than do present populations (Laws, 1977a). The amount of krill consumed by seals, birds, fish, and squid a century ago is unknown. There is evidence that penguin and seal populations have increased in .response to increasing krill availability as baleen whale populations decreased (Conroy, 1975; Laws, 1977b) . From an ecological standpoint it is reasonable to expect that the Antarctic marine ecosystem has adjusted to the changing whale populations, even though the adjustment may not yet have been completed. It is also reasonable to expect that other populations in the ecosystem will respond to changes in krill availability as harvest levels increase. D. Areas of Special Biological Importance The Scotia Sea is one of the regions of greatest krill abundance. Some fish species distributions are limited to the island groups of the Scotia Arc, Kerguelens, Crozet, Bouvet, South Georgia, the South Orkneys and the Antarctic peninsula. Areas of feeding and breeding for fish, birds, and mammals should be considered for protected areas as should migratory pathways for whales and other migratory populations. -34- E. History of Commercial Harvesting Although the southern ocean is remote, it is not un- touched by human activity. There is a long history of commercial harvesting there. 1. Whales Whaling occurred in the Antarctic in the 19th and 20th centuries. Modern whaling with factory ships began in 1904. Species of commercial interest have been blue, fin, sei, minke, humpback and sperm whales. Populations of all of these except minke whales have been greatly reduced in the last 30 years. Whaling is now regulated under the Inter- national Whaling Convention. Fin, blue, humpback and southern right whales are protected from harvest. Sei, minke, and sperm whales are harvested (Everson, 1977; Mackintosh and Brown 1974; Laws, 1977a). Blue whales are the largest of the whales, and conse- quently the most valuable to whalers and the earliest ex- ploited. As blue whales declined, the smaller fin, sei, and minke whales were fished. Changes in the proportions of species caught have been a result of reduction in abundance from exploitation, not from changing preferences. Until the 1960 's, the majority of world whaling occurred in the Antarctic. From 1945 to 1960 between 30,000 and 40,000 whales were caught in the Antarctic each year. Catches are now substantially reduced (Everson, 1977). 2. Seals Sealing has also taken place in the Antarctic. In the last century the sealers contributed some scientific infor- mation about the region (Deacon, 1977) . Except for a Norwegian exploratory expedition (0ritsland, 1970), pack ice seals have not been harvested for commercial purposes. Future harvesting of seals south of 60°S latitude will be regulated under the Convention for the Conservation of Antarctic Seals. In the early 19th century, fur seals were harvested until populations declined. Then elephant seals were harvested but harvesting became unprofitable before populations were reduced to a critical level. After shore based whaling was established at South Georgia, sealing again became economically feasible and some overexploitation may have occurred prior to 1953. There is now no commercial harvest of fur or elephant seals (Everson, 1977) . Harvesting is now regulated south of 60°S under the Convention for the Conservation of Antarctic Seals; no har V e.'7 -■■ r.g is taking place there at present. -35- 3. Fish Total reported catch in FAO statistical areas for the southern ocean (areas 48, 58, and 88) has been zero in recent years. However South Georgia and Kerguelen, known to be heavily fished, are included in reporting areas to the north (41 and 51) , Only southern poutassou and Patagonian hake are reported as separate species in FAO Yearbook of Fishery Statistics. It is probable that a large part of the catch of southern poutassou by the USSR in area 51 was taken in the Scotia Sea. Vessels of the USSR have fished near South Georgia and off Kerguelen (estimated catch 120,000 tons during the 1971-72 season). Principal species caught are Notothenia rossi, N. squamif rons, Channichthys rhinoceratus , and Champsocephalus gunnari (Appendix G. Everson, 1977) . Exploratory fishing has been done by West Germany, Japan, and Poland. Catches in area 41 which includes South Georgia built rapidly to 400,000 tons and then declined rapidly to a few thousand tons. Decline in total catch was due in part to a great reduction in catch rate (Everson, 1977) . Thus some fish stocks may already have been overexploited. 4. Krill Because of their great abundance and because of the de- cline in their major predators, baleen whales, krill have been receiving increasing international interest as a poten- tially harvestable resource in recent years. Exploratory fish- ing for krill has been carried out by the USSR, Japan, Chile, West Germany, Poland, Norway, Taiwan, East Germany, Spain, and Korea (Appendix F, Everson, 1977; Tetra Tech, 1978) . The most recent catch statistics available for krill (1974) indicate a total catch of 20-40,000 tons. The estimate of an average take of 200,000 tons per season in recent years (McWhinnie, 1974) represents about 0.3 - 1% of estimated stand- ing stock and about 0.3 - 0.8% of estimated annual production. Krill harvesting research has been carried out in the summer months, December though March, and has been concentrated in the Weddell, Scotia, and Ross Seas, and the East Wind Drift. Harvestable densities have been encountered most frequently around South Georgia, the South Orkney Islands, the South Shetland Islands, and the South Sandwich Islands (Tetra Tech, 1978) -36- The only krill species of cominercial interest at present is E_^ superba. Krill swarms are detected visually or acous- tically. Detection and catching methods are now sufficient to harvest more krill than can be processed with available faci- lities. The main obstacles to expansion of the fishery are now product development and marketing. A total catch on the order of a million tons is possible in the immediate future. Once a fishery of this size is established, it would tend to expand rapidly. Both scientific information on krill stocks and information on the effect of harvesting on krill stocks and on krill predators are needed (Everson, 1977). 5. Penguins Penguins have been harvested for their oil in the past. There is no present prospect for a commercial harvest of any Antarctic birds. Historically, Antarctic fisheries have developed around krill predators, thus harvesting the krill production indirectly. In the past commercial harvesting in Antarctic waters centered on seals and whales. Seals are no longer harvested, and the intensity of Antarctic whaling has been reduced in recent years. Commercial interest is now focused on fish and krill. Many nations are engaged in exploratory krill harvest- ing, and krill are expected to meet part of the increasing world demand for protein. Increasing fishing pressure on krill is anticipated in the next decade. F. Scientific Investigation A great deal of scientific investigation has been carried out in Antarctic waters, mainly since the turn of the century, e.g., the Discovery expeditions. Scientific activity has intensified in the years since the ratification of the Antarc- tic Treaty in accordance with the objective of peaceful uses for the region. Because the Antarctic marine region is relatively undis- turbed in comparison with other ocean areas, and because it is a polar environment, it is the subject of much scientific interest. Studies have been carried out on the continent and ice shelves, in open waters, in the pack ice region and on the islands. All the original signatories of the Antarctic Treaty, Argentina, Australia, Belgium, Chile, France, Japan, New Zealand, Norway, South Africa, the USSR, the United Kingdom, and the United States have participated, as well as Poland and West Germany. -37- G. Shipping Traffic There is considerable shipping traffic in the Drake Passage by vessels too large to pass through the Panama Canal. Commercial shipping, including oil tankers, passes through that area, usually north of the Antarctic Conver- gence which lies about midway between South America and the Antarctic Peninsula in the Drake Passage. In addition scientific vessels and commercial fishing vessels are found throughout the southern ocean at various times of year. Relatively few vessels penetrate the pack ice zone. H. Ecosystem Responses to Disturbance Even though the Antarctic marine ecosystem is relatively undisturbed in comparison with other oceanic regions, it is not a pristine area by any means. There has been harvesting of seals, whales, fish, krill, and penguins. Whaling has had the largest impact on stocks, with the populations of baleen whales drastically reduced from levels of a century ago. Because baleen whales consume enormous quantities of krill during the summer season, the reduction in whale numbers has reduced the impact of whale predation on krill populations and has allowed more krill to become available to other pre- dators. There is evidence that other populations within the ecosystem are adjusting to the changing abundance of whales. Penguin populations are increasing (Conroy, 1975) and there is evidence that seal populations are increasing also (Laws, 1977b). It is clear that the whale populations themselves are changing. Both population size and age at sexual maturity are different from original stock levels (Gambell, 1976; Laws, 1977a). The Antarctic marine ecosystem as well as the Antarctic continent have been used as a scientific laboratory. Data on the biology of the Antarctic marine ecosystem come from both scientific and commercial investigations. Carefully controlled harvesting with good data reporting can create an opportunity for a controlled experiment on the Antarctic marine ecosystem which can provide a great deal of information about its biology. I . Potentially Harvestable Resources An ecosystem viewpoint is a necessary perspective on the ecology and management of renewable resources in Antarctic waters. There are many potentially harvestable resources: fish, squid, seals, whales, krill and possibly birds. These populations are not independent but are closely related. -38- All of the others depend on krill as their main food supply. Krill are of central importance in the food web. Fluctuations in the size of krill populations will be reflected in the populations of its predators. In turn, fluctuations in the populations of predators, as exemplified by changes in whale populations, will affect the abundance of krill. At the other end of the food chain, diatoms are abundant south of the Convergence and there is a limited amount of benthic macro-algae. There is no apparent interest in har- vesting marine plants in the Antarctic at present. An ecosystem perspective on Antarctic marine living re- sources is feasible. Interrelationships among species have been identified and can be taken into account. Quantitative information is still required for most stocks and many preda- tion rates, but the structure of the ecosystem is known. J. No Native Peoples There are no indigenous human populations residing or using resources south of the Antarctic Convergence. -39- VI. ENVIRONMENTAL IMPACTS OF PROPOSED ACTION AND ALTERNATIVES The proposed federal action represents one point on a continuum of management regimes for Antarctic living marine resources, from totally unregulated harvesting to a ban on all commercial harvesting. Similarly the impacts of each of the alternatives under consideration also fall on a continuum from greatest to least impact on the marine environment. Future activities associated with the devel- opment of non-living resources in Antarctic waters could affect living resources through modifications of the en- vironment. For conservation of living resources, the activity of greatest concern in terms of environmental impact on the Antarctic marine ecosystem is commercial harvesting of renewable resources. Krill is the resource of most immediate interest, but there are other potentially harvestable resources as well: fish, squid, birds, and seals. Under different management regimes there would be different degrees of environmental impact. There are three classes of impacts of commercial har- vesting on the Antarctic marine ecosystem. The first and most significant are the impacts of harvesting on the popu- lations of the renewable resources. The impacts are both direct, i.e. changes in abundance of any stock which is harvested, and indirect, i.e. changes in abundance of species which depend on the harvested stock for food. The second are the impacts of shipping traffic and harvesting operations on the local marine environment. The third is the impact of increased human activity on a relatively un- disturbed marine ecosystem. The range of possible impacts of harvesting on stocks, on the marine environment, and on the southern ocean as a relatively undisturbed habitat will be discussed. Then the degrees of impact anticipated from the proposed federal action and the alternatives will be compared. A. Impacts of Commercial Harvesting 1. Impacts Unaffected by Proposed Action There are some environmental impacts of human activity in the Antarctic Ocean which will not be affected by the conclusion of the proposed conservation regime, specifical- ly sealing and whaling. Sei whales, minke whales and sperm whales are harvested in the Antarctic while fin, blue, hump- back and southern right whales are protected under the International Whaling Convention, Harvesting of crabeater. -40- leopard, Ross and Weddell seals is regulated under the Convention for the Conservation of Antarctic Seals, which has recently entered into force. The Convention for Antarctic Seals applies only south of 60°S latitude. All breeding by pack ice and fast ice seals does occur south of eCS although distributions of juveniles may occasional- ly extend north of that latitude during part of the year. Only those nations which are parties to the International Whaling Convention and the Convention for Conservation of Antarctic Seals are bound by the policies developed under those conventions. The possibility of unregulated har- vesting by other nations does exist. 2. Direct Impacts on Exploited Stocks The greatest environmental concerns over commercial harvesting in the Antarctic are the direct impacts of a harvest on the resource stocks themselves. The Scotia Sea, Weddell Sea region is the only known area of krill (E. superba) spawning. At the same time it is one of the areas of greatest abundance of krill and is likely to be a region subject to intensive harvesting. Overexploitation of gravid individuals before reproduction occurs would jeopardize maintenance of a breeding stock. The same problem may occur in other geographic regions of krill abundance, but that cannot be determined until it is known whether krill are one or several breeding stocks. Geographic considerations will be important in management of krill harvesting. Even though krill are very abundant, the possibility of their overexploitation is a real one. Krill abundance is estimated at 200 to 600 million tons in the summer. Since krill have a very long life span for zooplankton, their productivity may also be relatively low. The rate at which the krill population is replaced is not known at present. Therefore it i s impossible to estimate sustain- able yield with any accuracy. Nonetheless, expectations for krill harvesting are high. Present total world har- vest of marine resources is around 70 million metric tons per year. Estimates for potential annual krill harvest range up to 150 million tons in the scientific literature (Everson, 1°"1) and up to 200 million tons in the popular press (Gwynne, 1977). After consideration of possible re- placement rates, of increases in other krill predators with che decline of 'i.ales, and of the desirability of re- covery of baleen whaie populations, a sustainable annual harvest appears to be less than one-third of these esti- mates (Green, 1977) . A potential for an enthusiastic -41- unregulated harvest to overexploit the krill population is real . Commercial exploitation of krill may have direct im- pacts on the stock other than a change in overall popula- tion size. , Krill distribution patterns, productivity or secondary production rates, or behavior such as swarming patterns could also be affected. In evaluating the potential impacts of various har- vesting levels on krill populations, the different time scales of krill and predator populations must be taken into account. The life span of krill may be 2 to 4 years. The life span of krill predators such as a crabeater seal may be as much as 30 years. Since adjustments of predator populations to changes in krill abundance may take a long time, consumption by a large predator population v/hich has not yet adjusted to a decreasing krill population must be taken into account in estimating impacts of harvesting on the krill population as a whole. Time lags in response vary for different predators, and are factors to be con- sidered. The present level of krill harvesting in the explora- tory phase is roughly 20,000 - 40,000 tons per year, certainly too low for any direct impact on the krill stock. Practical considerations limit the total krill harvest now. Processing of krill must occur within a few hours of catch- ing, and processing capacity is limited. Markets are not yet extensive. However, with improving ability to locate swarms, increasing vessel and processing capacities, and rising demand for protein, krill harvesting operations may soon be profitable. Considering the potential need to redeploy long distance fishing fleets as a consequence of extension of national fisheries limits (Eddie, 1977), the potential capacity to overexploit krill probably exists already. Several fish species are already harvested in the Antarctic. Unregulated harvesting could lead rapidly to local overexploitation. The fishery techniques for fish harvesting are already well developed. The southern blue whiting, Micromesistius australis , is a migratory fish. It is present in the Antarctic only during the summer season while it is feeding there. The rest of the year it is in the Magellanic and New Zealand regions. Harvesting of this species would not be regula- ted while it is outside the Antarctic ecosystem under the -42- proposed conservation regime. The lack of regulation would be significant only in terms of the population of that one species. Should numbers of southern blue whiting decline, there would be at most a local impact on the rest of the Antarctic ecosystem. It could be replaced by other fish species in the diet of its predators. There is no present commercial interest in birds or cephalopods in the Antarctic, although that cannot be ruled out as future possibility. Hov/ever, no immediate direct impacts on populations of birds and squid are expected. Harvesting of Antarctic seals under the Convention for the Conservation of Antarctic Seals will only be regulated south of 60°S latitude. Under the proposed conservation regime, pelagic sealing could also be regulated between 60°S and the Convergence. Elephant and fur seals on islands near the Convergence are not now harvested. 3. Indirect Impacts on Dependent Species Concerns about the impacts of harvesting of one species on others which are dependent on it are most per- tinent to krill as the target species. Fish, squid, birds, seals and whales depend on krill as a main food item, and are in turn potential resources. Overexploitation of krill would be very dangerous to the Antarctic ecosystem. Consider the history of whaling. Most species of baleen whales have been reduced to roughly a tenth of their populations of a century ago. This change in whale numbers has caused some adjustments in other popu- lations in the Antarctic ecosystem, but because whales are the end of a food chain, the internal structure of the eco- system has remained essentially unchanged. However, should krill populations be reduced to a similar extent, enormous reductions in populations of fish, squid, seals and also whales would follow. Because of the crucial role of krill in the Antarctic marine food web, overexploitation would substantially alter the structure of the marine ecosystem and substantially damage its health. Even without overexploitation, a large krill harvest would affect populations of krill predators. Krill removed by harvesting would come out of the total now consumed by predators. With a decrease in available food, predator populations in intensively harvested areas would drop. Fin, blue, hiompback and minke whales are presently protected from harvest under the International Whaling •43- Convention. An increase in the populations of these species is desired. Because the age of first reproduc- tion for some of these species has decreased, a faster population recovery is possible. However, since baleen whales are entirely dependent on krill for food, a large krill harvest potentially would compete with whales for food supply. Probability of competition is increased since whale feeding and commercial harvesting would occur in the same geographic areas and at the same time of year. A reduction in food supply could cause a deterioration in physiological condition and in turn raise the age of first reproduction and slow the recovery rate of the populations. Further, a decrease in reproductive rates could contribute to a decrease in whale populations even without harvesting of whales. Crabeater seals are also likely to be affected by a krill harvest because they are very abundant and because their diet is predominately krill. Because of their abundance, crabeater seals are also the seal most likely to be considered for commercial harvest. The impact of a krill harvest should be mitigated by geographic separation. Krill harvesting is most likely to occur in open waters and crabeater seals inhabit the pack ice region. However, juvenile seels, whose survival is critical to overall seal population dynamics, may feed in open water or near the edge of the pack ice, competing directly with the krill fishery. Fish, birds and cephalopods which depend on krill as a major item of the food supply will be affected by a krill harvest. There is already evidence that bird populations are increasing due to increased krill abundance resulting from reduction in whale populations. Since harvesting will reduce krill availability, populations of predators are expected to decrease. Responses to a small krill harvest will be difficult to observe. Ecosystem interactions are complex. The direct and indirect impacts of harvesting on Antarctic populations as discussed above reflect the dependence of population size on food supply through a variety of mechanisms. Because the Antarctic marine ecosystem is dynamic and has already been under stress through harvesting of various populations, especially baleen whales, other kinds of sys- tem responses are possible. A change in the structure or character of the ecosystem such as a shift to a copepod- myctophid dominated food chain is possible. -44- 4. Local Impacts of Harvesting Operations The mechanics of commercial harvesting of krill will have local impacts on the marine environments. Krill are caught by trawling. The processing of krill can be done on the trawler acting autonomously, or may be carried out by mother processing vessels at sea, or may use shore based factory facilities. Because krill spoil within several hours after catching, processing must be relative- ly rapid. Trawling vessels will have to operate close to factory facilities. Most krill processing methods retain the soft flesh but discard the carapace of the organism. The hard parts of zooplankton are a normal component of detritus in sea water. In deep water, normal decomposition processes can handle krill parts which are dumped overboard. In shallow waters, such as around shore based processing facilities, discarded krill could accumulate, causing pollution prob- lems or perhaps attracting scavenging birds. Krill harvesting operations would also cause local pollution from ship bilges and garbage. Pollution prob- lems would be most pronounced in a shore based factory facility with shallow surrounding water and would be in- tensified if a large support staff were living in the vi- cinity during the harvesting season. At present, technological limitations on the rate of krill harvesting, limited market and availability of fishing vessels, have limited shipping traffic connected with krill harvesting. If krill harvesting is unregulated, shipping traffic will increase substantially within sev- eral decades, intensifying local pollution problems asso- ciated with harvesting and processing activities. Control on krill harvesting will incidentally regulate the level of shipping traffic. 5. Disturbance of the Marine Ecosystem The Antarctic Treaty and Agreed Measures provide for peaceful uses of the Antarctic continent, which have in- cluded a great deal of scientific study. These agree- ments do not apply to surrounding ocean. Even so, there is recognition that the Antarctic marine ecosystem is important for scientific investigation, because it is a relatively undisturbed marine ecosystem and because there is intrinsic interest in polar oceanography. -45- The Antarctic ecosystem is not untouched by human activity. Some harvesting of fish and seals and exten- sive harvesting of whales have already occurred. With increasing commercial harvesting of krill and possibly other species, there will be increasing disturbance of the marine ecosystem and the nature of the system will be increasingly farther from a "natural state" for scientific study. On the other hand, commercial data have provided information on stocks, abundance, reproductive and mortality rates and distributions, which otherwise would be unavailable. Data reported under the International Whaling Convention are an example. Such information cannot readily be collected on strictly scientific cruises. From a scientific standpoint, a carefully controlled har- vest of various species with good monitoring and data reporting could provide an opportunity for a controlled experiment on the Antarctic marine ecosystem. Much could be learned about responses of the system to different levels of harvesting. In addition, such a controlled ex- periment could provide data which are needed for effec- tive conservation of Antarctic living resources. More information is necessary to be able to manage for long term productivity of the ecosystem. Some increase in the impact of human activities on the Antarctic marine environment appears to be inevitable. Commercial interest in krill harvesting is already develop- ing, and harvesting levels are likely to increase as world protein demand rises. An increase in harvesting south of the Antarctic Convergence is expected even though the U.S. has no immediate plans to participate in it. Harvesting will increase whether a conservation regime is negotiated or not and regardless of whether the U.S. is a party to it. The purpose of a conservation regime and the motivation for U.S. participation in one is the mitigation of the impacts of uncontrolled harvesting. B. Environmental Impacts of Proposed Federal Action and Alternatives The approaches to negotiating a conservation regime for Antarctic marine living resources, as indicated in the proposed federal action and alternatives to it, comprise six alternatives identified as (1) no action on a conserva- tion regime, which implies no conservation measures, (2) individual national action on conservation policies and some coordination between these policies, (3) agreement on -46- a regime for collection and transmittal of data at present, but postponing commitment to a conservation mechanism until later, (4) negotiation of a complete conservation regime providing for both data collection and an organization to establish conservation measures and regulate harvesting, (5) negotiation of a complete conservation regime, plus the establishment of quotas for harvesting of all resources or establishment of a moratorium on harvesting, and (6) prohi- bition of all harvesting in the Antarctic region. The feasibility of negotiating each alternative, the ability to develop and enforce conservation measures , and level of harvesting activities anticipated under each alternative are presented in Table 2. The probabilities of overexploiting the various resource populations under each alternative are indicated in Table 3. Indirect im- pacts on stocks are compared in Table 4. Anticipated im- pacts on local marine environments and on the health of the ecosystem are indicated in Table 5. The effects of the alternative approaches on science are compared in Table 6. 1. Ban on Harvesting The sixth alternative, prohibition of all harvesting, would imply no harvesting activities south of the Antarctic Convergence, if such a regime could be negotiated and en- forced. This alternative would provide for the least impact on the Antarctic marine environment. It is the most favor- able alternative for recovery of whale stocks. Small ad- justments in other population sizes would be expected as whale populations increased. A ban on harvesting would preclude the possibility of overexploitation of any stocks and would eliminate direct and indirect impacts of harvesting. Since seals and whales could be harvested under the International Whaling Convention and the Convention for Conservation of Antarctic Seals, small adjustments to other populations in response to sealing and whaling could be expected. With the ban on harvesting, no commercial data would be available, thus eliminating a significant source of infor- mation on the stocks of some populations. Even though this alternative represents the least envi- ronmental impact for the Antarctic marine ecosystem, achieve- ment of a total ban on all harvesting is an unrealistic expectation. Such an agreement would not have the support of nations presently involved in exploratory krill harvesting, -47- so that a total cessation of all harvesting could not be achieved. 2. No Action The contrasting extreme of possible alternative action is no effort to negotiate a conservation regime and conse- quently no conservation measures for Antarctic waters. The consequences of this alternative are unrestricted and un- regulated harvesting of all potentially commercial species. In time, technical and economic limitations on the harvest- ing of krill will be overcome. Intensive harvesting can be expected over the next several decades. An unregulated har- vest will result in overexploitation of all harvested species This alternative represents the greatest impact on the stocks of renewable resources. Direct impacts on the sepa- rate stocks as they are harvested can be expected from over- exploitation. In addition, indirect impacts on seals, whales fish, birds and cephalopods from exploitation of krill would be maximum. With overexploitation of krill or another species, the balance within the Antarctic system and the structure of trophic relationships can be upset, thus seriously damaging the health of the ecosystem. Unregulated harvesting will operate to maximize short term gains, thus potentially decreasing the overall productivity of the Antarctic eco- system over the next century. This alternative represents the greatest possibility of local pollution problems from harvesting activities. In the absence of an agreement it is expected that commercial data will not be reported completely so that this source of information will be lost. This alternative would result in the greatest deviation from a natural or undisturbed state for the Antarctic ecosystem. The actions of the U.S. and other Treaty nations at the Ninth Consultative Meeting indicate that they do not support this alternative (see resolution IX. 2. in Appendix A) . However it is the default alternative should negotia- tions fail to establish a regime. 3. National Action The second alternative is separate action on conser- vation of Antarctic living resources by nations interested in harvesting and some informal coordination between pol- icies. Lacking consistent management objectives and a procedure for establishing and enforcing conservation -48- ■u C 0) e a O TJ .-1 c 0) n3 > %C ^ :3 •H i-H W •H OJ ^ !-i •H 3 C/] Cfl CO CO CU OJ 14-1 e - c c/2 O C -H O 4-) •H CO 4-1 > CO )-i U CD (U c/3 a c o o o M C'^-i •H O •U cn 4-1 (U U E CO CO (U OJ 4:: u > u •H M-l 0 4-1 0 M-l CO C c .-H QJ V^ (U cu >T3 4-) OJ C r-l hJ CO CO QJ 0 0 0 5-1 ^1 Z S ^-1 CU 3 0 en CO 14-1 C CO c 0 QJ W cj e C 0 •H 4-1 CO CO a > QJ 0 0 0 j-i !-4 S S ,-H QJ 3 QJ cn CO > c CO QJ 0 QJ Q 0 E >. :>. 1 CO QJ >s CO 4-1 t3 !-i 13 •H > r-l CO QJ 13 1 ^ •H QJ -H QJ •H .-1 >. QJ > QJ QJ QJ CO.-I N CO 4-1 4-1 CO 5-1 CO QJ 4-1 13 U 0 •H •H QJ :>> CO . J-l 4-1 CO •H CO 3 CO •r4 to 0 bO •H^ c -a ^ QJ CO QJ .-1 C 4-1 4J 4:: p,^:: QJ CO E q E 4-) •H txO c U QJ > d w C C 4-1 CO 0 5-1 QJ •iH QJ •H CO 0 Id QJ QJ 4-1 QJ CO QJ QJ 3 x: CO QJ 4J QJ 13 4J 1—1 CO 0 1-1 4-1 CO 13 M-l 4-1 CO I-l 13 5-4 4J CO 5-1 q CO 0 QJ QJ CO rO C 4-1 0 QJ >-l CO 0 c QJ 0 q 4J •1-4 -H >^ PL,IW Pi CO CO •H CO ffi T3 CO w > CO 0.4-) M to to 43 CO C T3 QJ TJ •H 1 ■ C 13 4-1 1 13 • 1 1 bO QJ QJ > QJ CO CO CO CO QJ CO 1 u 5-1 5-1 CD 4-1 - 4-1 •rH > QJ QJ 4-) 4-1 0 3 CO QJ QJ 4-1 •rH QJ t3 0 4-1 •H T3 x: > •H •H -H txO^ > 13 13 q e N C !-i 0 CO QJ CO yi 0 E e QJ QJ .— 1 -I-l CO E ■H -H CO a CO C 4-1 r-l CO <—* •rH 0 !-i CO :3 3 to > 5-1 13 1—1 CO QJ U :-l to JC a I-l c CO 0 0 q 13 QJ QJ CO CO .— t 4-1 -H CO !-i X 0 13 X :s 0 CO 4-1 4-1 >.-u a< ,— 1 C U CO T3 QJ >^ U QJ bO 0 CO t—l QJ •H CO •H 4J M-( QJ QJ M r-H QJ 4-1 C CO E M P- ^ QJ ^ CO 0 >.4-l QJ 1—1 •H •H • QJ E u q •H CO 1-H CO - QJ QJ CO > CO 13 E 4-1 to -H QJ QJ 0 0 ■r-( 14-1 CO CO !-l 5-1 bflO nH 0 •H C •H 5-1 QJ 4J 4-) 4-> r-l •H 4-1 QJ QJ 0 C C u^ 3 4-1 CO 1— 1 0 to •H CO 4-1 •H - •i-l CO E P -H 1 W) c •H (X CD 5-1 >^4J 4:: C C >^ CO 4-1 CO 4-1 0 QJ •iH q >^ QJ QJ QJ 0 to 5-1 bO < •H bD U •H QJ QJ - CO .—1 !-i •H (1013 5-1 5-4 -H 0 0 -M 0 •H ,— 1 U E 13 QJ C 4-1 0 •r-l 0 5-1 U 4:: QJ • CO bOr-l E •H 0 0 QJ > c :3 r-H bUi-i > 13 q CX QJ CO :s 4J QJ c 0 0 0 G U 4-J )-i •H 3 P 0 0 q •I-l 4-1 q •H •H C c CO •r4 QJ CO CO CO CO •H q !-i CD 1— 1 QJ 1 3 QJ 4-1 4-1 ^ 0 <+-! ^ .-1 4n CO QJ QJ 4-1 4:: a bO CO 13 0 E •rH CO u 0 T3 in QJ E !-i ^ CO 0 q C C-r^ to QJ > QJ QJ 0 QJ C CO bO >s'H d QJ QJ 13 0 •H 0 5-1 q 13 bO •H > 4-J !-i CO QJ QJ > U CO .-1 c > 4-1 1— 1 •H 4-1 tH 5-1 0 I-l CO •P !-i •T3 0 E •H J-< CO QJ CO I-l 0 5-1 3 4-1 CO 4-1 QJ •H q q a CO >% C .C QJ 4-1 C QJ Ci,< •H -H CO >. 0 CO QJ CD > 4-1 0 CD < ffi rQ CO CO T3 •H ;=i 4:; CO ^"^ [5 4J ■X J^ ^ i-H > c 0 CD :§ E CO c QJ 0 > ■~ 1 13 •I-l •H C 5-1 C 4-1 4-1 0 QJ CO 1—1 CD CO CO •iH CO QJ CO c c i-i C C !-i C C •r-f V4 0 0 0 :i 0 0 13 QJ CO 0 •t-l CO •H -H !-i 4-) 4-1 CO 4-1 4-1 0 r-l 0 0 CO QJ CO 0 0 < Z C > E S CO 0 -49- X) QJ . n t3 ^ W OJ o a. • QJ C rt M CO CO C Xi (U (U (U 4-J M 4-1 >^ >^ 4-1 O -H Z 13 QJ • . .. X) [3 CO QJ O Ph • Q) C CO !-i CO CO P rC 0) Q) QJ 4-15-14-1 >^ >H 4-) O OJ CO O S P-.'-H S — . 1 ~l XJ O 1 1 1 en 1 en 1 -H QJ 4-) 4-1 p u QJ -H 4-J 1 1 to > CO •H QJ 4-1 5-1 4-1 P •rl QJ 4-1 ,-t >, - <\) X) P E •H O X) QJ rH QJ QJ >. o en 5 > CJ c 0) CO :3 bo rH E QJ 5h box ?> QJ • cd O •H •H E •r-l n QJ . cr QJ E ' m bo P d^ bO p tj en Q) C M-l E -C QJ c CO rH QJ QJ C d 4J O C o bO^H CO O 4-1 -H 5h Q) O O Q) CO X > 5-1 •H QJ •H • 3 5h rH 4-1 n P •rl en (U c CO 5-1 rC en CO •rl r-l 5^ b04-l 4J en CO Mh QJ CO O 4-1 CX > J-i O 4-1 tsO O 5-1 C 4-1 X 13 • O 5h en C en QJ O >. E P tn •H CO 0) cu O CO 03 QJ CO -H O bO<4-l C 4-1 4-1 O QJ O CO Q) o CO QJ O Q) > ^ -H bO QJ QJ E QJ CO QJ 3 QJ O CO- a > •H 4-1 cn P T3 4-1 12 QJ •H 5h •H !-l rC C 5-1 t:Xl a-u •r-, O 5-1 -H u c a VI 4-1 o p Xl O P 5h 4-1 4-1 QJ CJ CO T3 QJ P c a.4-1 x j::: CO 4-1 5 ° p CO CO p CO QJ O QJ O bO to C O (X, CO 13 QJ 4-1 •H O < CO O 4-1 O CO S & en x: P o- o ShX E z to p •rl to 1 1 QJ 4-1 4-1 E 5h O 4-J QJ 0) rH O en 1 Q) QJ O >i > 1 4-1 c • • QJ r-l •H •H [5 O >, en CO C 5-1 bO CO 15 E 5-1 rH 4J O bO O P bO CO Q) QJ QJ •rl^ CO d O rH C ,-{ O iJ •H .-H 5-1 > 5-1 5-1 en 5h x: 14H CO P. O QJ 5h rH O 4-1 CO O O O 4-1 M-l CO Q) X rH X QJ O D •rl U M-l QJ cn O QJ rH 5h 4-J QJ x; C cr > •ri CO 5-1 >^ QJ >. CO CO CO 5h 5h en 4-1 XI •rl tio en ^ O TJ hO > >. 4-1 O QJ 0) O 4-1 •rl O bO 4-) O C CO M-i QJ O 5-1 4-1 in o > > Mh en QJ QJ C O r^ O -H QJ 4-1 rH CO •rl ^ 4-1 en 5h O QJ 4J •rl CO o -H pt; xi CO o x; rH QJ CO QJ > 1 4-1 C JJ •H c •rl QJ C •rl X o bO . 5h :>.4-i bO ^ CO c 4-1 X; C1.X1 4-1 QJ O c CO >. CO rH X QJ C O 4-1 • O O O QJ •rl CO x: QJ C 4-1 X: QJ en •rl QJ •r^ en •H bO QJ QJ en -rl H cu ■ 4-1 CO •H >'T3 4-) 4-1 E M 4-) QJ 4-1 ^ en 'T3 en TD d E > QJ QJ QJ MH CO •i^ CO CO C O QJ QJ X •rl 4-1 rH 4J O QJ C ^ 0) 4-1 c o 0-. E • r—^ 4-1 4-1 CO •H > O >,tA QJ O 4-1 E QJ rH CO - C (J •H 4-1 E 5h • >H ^~^ o O XI 3 -rl 5h CO rH en o d TJ O •H QJ bO to 'O >,-H O <-! >.-ui o d d QJ •H 'T3 QJ CO rH bO C Xi CO rH ^-N 4-1 •H 5h 4-1 •rl 4-1 E C -u O QJ 5-1 QJ O CO C QJ QJ rH CO O CO X en (U QJ -H QJ 5-4 rH QJ E a 5h O rH E rO CO 4-J QJ •H QJ ,— 1 QJ CO E E QJ (X en •H O Q) -H (X-r^ CO 4-J CO > x; > ^ 5-1 -u E 4-1 E C bO 5h t3 -U E bD4-i O 5h 5h o u nj tiO CO O o CO O O Q) P-i QJ U O QJ cn p O CO 5h CO H <: 13 a ■U r-l O CJ J^ v-'l4H CO O 5h QJ cr E -i: PL, X u 0) •D c 3 m C O •H 4-1 td i-i :3 ex o a (U o M ::! o m (U n OD ;3 o CO •iH ct •H > ;-) CO OJ C x. M 4-J 0) 4-1 U-l .-1 o CD C T3 O C •H ctl 4-) CO C +-J O •H •H O •U I— 1 a (X CO X QJ I— 1 5-1 CO 0) V4 > 0) O T3 (U M-l M-l O X) >. QJ i-> CO •H o i-< a •H o ^ u •H tx CO CO 0) O ^ P-i ■Ul OO 0) I-I rO CO H -so- c q q O o o c •H C •rH q •H •H C/3 CO •i-l CO CO •H CO to -o g 1— 1 •H 'T3 ls ^ •H T3 15 rH -rl , CO g QJ O >^ CO g QJ O >. to g U C X^ C E 4-1 c ^ c g 4-1 q XI q E •H O O •iH o O •r-l o o CO O - ^3 -rH o O -T3 -rH U O -13 •H CJ (U I— t 4-) QJ 4-) 1—1 4J QJ 4-1 r-H 4-1 QJ 4-1 .—1 D- CO 4-1 Cfl bO CX CO 4-1 CO W) CX CO 4-1 CO bO CO X CO CO c C X CO CO C q X CO CO q q r; 01 D-i-i !-i •H QJ CXrH U •H QJ CXr-{ 5h -H 3 !-i P QJ .-^ >-< P QJ rH !h P QJ rH QJ QJ OJJ-P CO QJ QJ b04J cfl QJ QJ 50 -U CO > 4= QJ C ^ > JH QJ C r; > X QJ !=:£ O 4J J-i M 3 O 4J 5h M 3 O 4J 5h M 3 ^ M-l CO ^ MH CO rC Mh CO ^ !-i O .-H 4-1 5H O rH . 4-) 5h O rH P M O CO P 5h O tfl P V4 O CO O QJ M-l C QJ O QJ Mh q QJ O QJ Mh q QJ CO 4J O zn CO 4-1 O C/J CO 4-) o w M-4 C •H MH C •iH Mh q -H •-T3 CO O 4-1 o ■ -. T3 CO O 4-1 O ■-13 CO O 4-1 o CO >^ q; •H Cl! •H >, QJ -H CO •rH >. QJ •H CO -H .—1 >-H 4J Cn 4-1 > 4-) r— 1 XJ en 4-) > 4-) rH 4-1 CO 4-J > 4-) CO QJ CTJO q !-i U QJ COO c 5h O QJ COO q 5h o 0) ^ r-H O QJ QJ 5-1 ,^ 1-1 O QJ QJ 5h ^ 1-1 O QJ QJ 5h cn •H 3 ^ > CO cU •H p vO > CO CO •rH P ^ > CO CO i-H W) C C 4-) 1—1 hO q q 4-1 r-t bO q q 4-1 C QJ M-l O O C C QJ Mh O o q q QJ Mh o o q tD U O O O < 1=1 5h O U o < ;=) 5h O O CJ < ^^ 5h 1 x: 1 o 1 O 0) CO ^ T3 U M-l g-d 5h mh ;5 13 x ^ -H -> o a QJ o q QJ o q O --Mh CO CO t-H CO to 4-1 >. 1—1 CO CO 4-J >^ rH CO CO CO 4-1 -o-o 13 M-l C t— 1 13) Mh q 1— I 13 rH CO 5h U O >. CO O O •iH QJ >^ CO O O •H QJ >^ CO O QJ O •iH CX .-1 T3 fX ^ 1-1 -a CX ^ :-^ 13 P.MH U Mh pq o 4J S-l O QJ M-l •iH U )H O QJ Mh •H 4-1 5h O O QJ ,—1 C -H ,-1 c/) O rH c •H i-H CO O rH q -H ,-H XI >. - CO QJ ^ CO p QJ ^ CO p QJ X CO QJ q rH x: x: CO 4=: cfl^ ■'x: CO ^ CO ^ ■-x: CO JC to -H QJ CO (X QJ U ex a o ■u CO QJ 5h (X U o 4J CO QJ Sh CX p ^ •H 0) U O QJ QJ CO CO -iH 5-1 O QJ QJ CO CO -H 5h O QJ CO Mh -H [J^ CJ fX<4-l o ^ 1—1 QJ M-l CXM-i a rCi rH QJ Mh CXMh U O O rH 1 CO 1 CO OJ 1 !h Q) .. QJ QJ QJ QJ •- O QJ e QJ X) g QJ -13 g QJ Mh X3 E !h >. CT3 g !h >^ CO E 5-1 QJ 13 ,-H •H m 1— 1 O •H P i-H O •rH P X q QJ rH 4-1 QJ QJ 4-J QJ QJ 4-1 to x: •H C lU ^ -d c P^T^ q P QJ o to U •H M-l •H •H MH -H •H Mh rH •H t^ --I C .-< q X QJ 1—1 ,-( QJ •H t— 1 QJ -iH rH QJ -rH E rO i-H 4-) >.^ rH 4-> >^rC rH 4J CO •H to CD CO U 4-J CO CO 5h 4-1 CO CO to bD4-l E-H Q) -f-l 6 •H QJ -H g -rH O QJ CO c/2 T3 > s CO 13 > [S CO 13 (X 5h QJ CO q O - 4-J ■- 1 13 -rH O 4-1 •rA C 5-1 q XI XI q q E 4-1 O QJ CO i-H tT3 CO q 0) CO CO •H CO QJ cfi q QJ CO g •H C J-J q C V4 d q -H E X) 4-) q ^ O O O P o O T3 QJ CO -iH 03 5h QJ CO O •H CO •H •H ^4 QJ 13 g x: QJ U 4-1 CO 4J 4-1 o 5h g a 4-1 rH o o CO QJ CO u o txo q o QJ to < 2 C > e 2 CO o i 03 CU o >^03 QJ O >, 03 ■U C 43 C 4-1 C XI C 4-1 q XI P •H o C •H O c •H O C O ■- 13 -H O o ■- X) -H o O •-T3-r4 O 1— 1 4-J (U 4-1 •H i-H 4-1 QJ 4-1 •iH r-l 4-) QJ 4-1 -H fX W 4-1 03 0£ 1 CO (X CO 4-1 0) 00 CO Ph CO 4-J CT3 00 CO X 03 03 C c CO >< 03 03 C c CO X 03 03 C C CO (U CLrH !-( •H •H 0) (X.-I 5^ •rH •r-l QJ CXi-l 5-1 -H -H !-i P OJ ^ g 5-1 P QJ r-l p 5-1 P QJ .-1 g 0) 0) 00 4-1 03 g OJ QJ 00 4-1 OJ g QJ QJ 004-1 03 g >^ CU c g o > ^ QJ C r; o > X QJ C X O O 4-1 5-1 M CLJ O 4-1 5-1 M CJ) O 4-1 5-1 M 3 O 5-1 C 1 5-1 5-1 c 1 Vl ^1 C 5-1 •-X) ^ 0) M-4 4-) •H CO O •H >^ CU M-l 4J •rH CO O •H >. QJ M-l 4-1 tH CO o •r4 i-H 4-1 O M-l 4-1 C 4-1 I— 1 4-1 O M-l 4J c 4-1 ,-H 4-J O M-l 4J C 4-1 Qj re 03 C O c O CU 03 03 C o C O QJ 03 03 q O c O ^.-1 x; •r-l 03 I— 1 •H p 4-1 cn > •H 03 j—< •r-l P 4-1 CO > •rH 03 rH .-1 bO :30 C 5-1 4J 4-1 03 i-l 00 po c 5-1 4-1 4-J 03 r-l 00 PO C 5-1 4-1 4J n3 C oj O O o O 03 C QJ C OJ O O o O 03 C 01 C QJ O O O O 03 C QJ 1=1 M W MD C_) M-l > < en 1=1 5-1 CO ^ c_; M-l Xcn P 5-1 CO vO UM-4 > < CO ^ 1 00 -C 1 00 m .—1 c CO t-H C •H CC ;5 01 o •rH 03 15 QJ o U-i o 4-1 >-H M-4 O 4-1 --H CO 1 t— 1 03 CO 1 ,— 1 0) ■-I c 5-1 •- •H t3 ■-1 C 5-1 ■- •H T3 cfl O 0) t;) 0) T3 c 03 O QJ t3 QJ TD c O -r-l > 0) W CU 03 O -rH > 01 CO QJ Oi O 4-) O 4J •H g O 4-1 O 4J •H g I— 1 03 •H [5 g OJ r-l 03 •H ^ g QJ >— 1 >. O 5-1 •H 5-1 I— 1 f^ O 5-1 •iH 5-1 QJ ^3 X) :-! 0) :3 g 01 p T3 .-1 QJ P g QJ E a 03 (x^ 5-1 4-) 5-J B ex 03 (XrC 5-1 4-1 5-1 C o o 0) X 4J O P 0) O O QJ >^ 4-) O P QJ O W ex 5-1 CU O <+-l M-l 4J CO (X 5-1 CU O M-l M-l 4-1 z •H •H T3 X) QJ 5-1 CU 5-1 g - -. Q) g * "^ QJ B g CU > •H M o B •H ^1 O g d 5-1 P 5-1 C 4J nH CU C 4-1 .-1 0) ^^ •H :3 03 4-1 •H p 03 4-) -o M-l B M-l g 0) OJ •H 00 0) •H 00 QJ ^1 C 0) c c C o; c c C C O 4-1 •H O O 4-1 •H O O •H 4-J 2 03 6-H S 03 gr-l s o 00 u C C r-l s^ O X! g •H 4-1 r-l 03 00 4-1 -o c x: u P CO a • 0) 03 (U Q) 03 CO O •H QJ 4J •r^ CO 4-1 > W 1 r-( . ---N 4-1 •H 5-1 > •i-l 4-) ^ e C 00 5-n3 4-1 g 004-1 O 5-1 O ^1 CO o o U to CO +J 0) C > 0) •H +J +J o CO cu c S-i M-l (U O 4J I— 1 a; CO u C X) CO C T3 CO C P C rO O CO •H 4-) C O •H CO cn >— 1 (U CO bO 5-1 C (U CO -Tj rC dJ U LM <]■' . •H p O cu >. •rH 3 O CU >. •rH P O b ^-^ ■- O -iH ^ > ti 1— t •- O tH ^ > R i-l ■- O •r^ •H o •H d QJ p CO •H o •H a •H 4-1 CO c: 5^ ^ CO > CU 1-4 -4 5-1 c; 54.-H -a p O -H bO o ^ CU 5-1 c: O CO TJ ^ -f-i M-4 i-H 4J •H CO [5 CO CU CO >% CO > CU i-H CU 5-1 O CX U e •H CO 5 CO CU CO >-, CO > CU I-H CU 5h U CX CJ CU CO p CX C CO 4n 13 p CO CO CU CO 13 CU 1-4 5h C 5h I-H 13 p CJ -H bO O rCi , CO > (U I-H CU 5h CJ CX O CU CO P CX C CO ^ 13 P CO CO CU CO 13 (U rH 5h C 5h I-H 13 7i O -r^ bO O JO -i CO > CU I-H CU 5-1 O CX CJ 0) CO p CX C CO j:: 13 p CO CO CU CO 13 CU rH 5h C 5h rH 13 p U -iH bO O JD CU 5-1 C O CO 'O a; -iH Mh I-H 4-1 •H CO 15 CO (U CO >^ CO > Cm w CU CO CO 4J CX QJ •H -i CO o CO >H CO 0 >^ g D- c a E (xc a rH 0 c o 0 QJ 4-) 0 i QJ UH •H fx, ^ •rl r-H O 4-1 M rH 0 4-1 >-i •rl }H 4-J ^ CO CO QJ rH > > rH ^ O 13 O XI U 3 0 4-1 0 P O C Oh O 0 C Ph 0 CO 0 CX }-i CO o cu )H CO 0 QJ 0 QJ 0 a c ex )-i ex C D- !h E u (X T3 X -O X CO o 4-1 0 4-) C C 1 r-l 1 O •rl rH 1 0 •rl 0 •H C QJ r-( 4-) M-l " CO ;s rH 4J 14H « CO 5 •rl •rH rH •H CO >. , CO xt •H CO >^ CO 13 4-) QJ CO Is en 0) CO 4-1 QJ c QJ ^ CO QJ CO 4-1 QJ C QJ CO CO 0 X CO > QJ •H r-l CO rH CO > QJ •H ,—t CO rH rH CO 4J 3: C U <-i •H C 0 > QJ (X, CO XI :3 o -H bo CO 4-1 CO 0 5-1 3 0 -H bO CO 4J CO a U E >^ CO QJ S rO -i C > PX c CO XI Q) Jh n > PX c CO 0 CO QJ Sh p CO T3 ^ -H CO XI CJ :3x CO T3 ^ •H CO X 0 PX CO E }h 0 C -r) XI 'V X CO o 4-J 0 4-1 C C 1 r-l 1 O •rl rH 1 0 •rH 0 •rl C QJ r-l 4-14-1 - CO [5 rH 4J <4H - CO 13 •H •H rH •H CO >^ CO T3 •rl CO >^ CO 13 4-1 QJ CO IS CO 0) •H rH CO rH CO > QJ •rl rH CO <—{ ,-\ CO 4-1 [5 CU !^ CJ rH X r-l QJ Sh U .-H XrH P QJ O 0) CO Id •H X 4-1 0 0 QJ CO p •rlX 4-1 0 a u QJ bO C CO 4:; 73 XI 0 5-1 4-1 C CO X T3 X CJ !h 4-1 0 0 CO ci; CO CO CO (U CO 3 QJ 4J CO CO CO QJ CO P QJ 4J CO (X, QJ C -t-l Sh ^O QJ rH S-l r-l e bO C QJ TJ QJ rH !-l rH E bO C QJ 13 0 CO QJ q ^^r-l •H C 0 > C V4 rH •H C 0 > QJ CX CO X3 :3 0 -rl M CO 4J CO 0 !-i p 0 •H bO CO 4-1 CO U M E >^ CO QJ E XI CU V4 C > px; c CO X QJ !h C > PX c CO 0 CO QJ Vh p CO X) ^ -H CO X 0 DX CO "13 ,ili •rl CO X U I3X CO E Sh 0 C3 ^4 1 U 1 QJ CO c 1 CO c 1 rH 4J 0) ^Xi 6 ---H 4-1 CO XI •rl t-\ 4-1 E --T^ 4-1 COX •rl ,-t -U 13 0 pi d rH CO CO QJ (U 4-J CO QJ 0 & 0 CO CO QJ QJ 4-) CO Q) 0 [3 0 CO CO 4J •H CO X > y~\ bO 5-1 0 QJ CO !h QJ bO !h 0 QJ CO U QJ CO 13 CO Sh X) >^ C ^) :3 > CO CO c 4-J > >.C P P > CO CO C 4-1 > QJ U CO d XI QJ (U J-i CO 4-1 T3 5-1 QJ CO CO c n V-i CO 4-1 13 !h QJ CO CO C U E C QJ p 0 CO 2 0) x: ;3 QJ CO ^1 QJ X 0 CO QJ XI P QJ CO U QJ X 0 CO 0 QJ Sh X Sh XI •H 4-) > 04-1 M XI CJ rH 4-1 0 X > 0 Mh 5-1 X 0^ 4-1 OX CO E 0 CO CX 0 o c « u 0 1 v^ •rl 4J Td 13 c x: ^^ § 4-1 CO bO " QJ CO 0) QJ CO CO 0 • •rl QJ 4J •rl CO r-l , /-s 4-1 tH M > •rl 4-1 QJ !-l QJ 0 CO c QJ QJ rH CO 0 u XI CO OJ r-l QJ E CX !-l 0 rH E XI CO 4-1 CO •rl QJ r-t ex CO -rl 0 QJ •r4 CX-H CO 4-1 CO X XI > rd g C M 5-1 Xl 4-1 E b04-i 0 u bO 0 rH Sh CO 0 0 0) PL, QJ 0 0 QJ CO 0 0 C C U rH CO H a a 5-1^-^14-1 CO u )H QJ cr E 0 •(-I P-l CO X -54- JH 4-) IM O >^ ■U •H > •H 4-> O D TD O VI Oh t3 en C (U CT3 > •H x: J-J W CO I— 1 c CO 5-1 OJ (U -C 4-1 I— 1 0) CO ^ 4-) X) C X) CO c: CO C O j-j •H c 4-) (U O e CO c O X) u QJ •H cn > o c a . CD cn a O e CJ I— 1 0) in 0) r^ ,Q nJ H >. >^ QJ rH 1—1 1 ec 1 0 1—1 1 0 1-1 en CO c--- • en CO C -^ • en CO 0 QJ . en ^ C 4-1 XI QJ ^ C 4J x QJ S •r4 Q) CO V-l 4-J 0 -H CO >^ en 0 •H CO >. en 0 > en - — 0 0 C cn .-1 QJ i-l >-l c en 1—1 QJ 1—1 Sh C QJ X) 1-H 0.4-1 CO CO :3 S-I QJ QJ CO 3 S-I QJ 0) 1—1 CO QJ •H CJ (X g QJ bO 0 ^ > & Q) bD 0 X > 5 X > 4:: CO E 0 !-i QJX •H X) 0 S-I QJ x; -1-1 73 0 •H 0 QJ CO U-l -H J 0 J-i CO .-1 CO hJ 0 5-1 CO 1-1 CO hJ cn 4J 1-H QJ 1 1 !-i 1 1 4-1 4-1 1 1 4J 4J - 3 c -o cn cn Q) C x) cn en QJ E c 1 c X. 4-) 0 CO >-. QJ cn 0 CO >^ QJ cn •H 0 en 73 0 4-) 0 •H M cn > {X •H 5-1 cn > CX 5h •H QJ CO •H ■— t :3 4J QJ 5-1 :i 4-1 QJ S-I P QJ 4-1 -H QJ 4J CO u e CO c bOU-i CO 5-1 CO C bC+A CO 5h 4-1 CO 0 5h c; CO > r-4 ,-H U >-l > 0 . 1-1 1-1 0 S-I > 0 QJ 1-H :i 4-1 0 •H > en D-,r-l 3 0 en QJ CLn-l Id 0 cn QJ 1-H a QJ CO c CX CO !-i 0 X -H >^-Ul M-l QJ 0 X •H >^4-l mh QJ 0 X X E 0 QJ 4-1 QJ CJ QJ J-l ,-1 0 0 •H c; QJ S-I i-H 0 0 •H C •iH QJ 0 E Sh cn C en 0) !-i ^ ^ 13 . a CO 5-4^ 1-1 D • CJ CO cn Sh en 1-H 4J QJ QJ •H QJ QJ CO S-I E bO QJ 1-1 QJ CO S-I S 50 QJ 1-H en QJ 1-H cn x; N CO !-l M-< > M-l 0 4-1 QJ C ex CO > <4-l a 4J QJ C CX CO 0 >'4H •H P 4-1 -H s a. 0 0 0 •H en 4-1 -iH en x 0 0 -H en 4J •H en X Ph 0 0 :3 •1—1 0 en >> QJ QJ Mh 1 4-1 4J 4-1 .-1 a 1 4J 1— 1 U 1 E 0 0) CJ •d 6 5-1 r-l 3 4-1 S-I 1-1 P 4-1 •H 0 d > QJ • 0 •H 13 a 0 -1-1 13 0 5h bO ;3 x) hO-i-l 4J >,rC 5 QJ d x: ^ QJ :3 QJ C:: T3 0 c 4-) en 1-H cn !^ T3 cn 5-1 x! 4-1 •H QJ 5h 0 0 >^ QJ cn 0 en 0 C 4-) Sh CX .— 1 :i cn ^ bOTJ >. S-i bOTJ >^ S-I •H en TJ •H C 1-1 t—i a . C i-H 1-1 CL QJ C E a 0 QJ .—1 -iH QJ i-l >>-i-l QJ 1-1 QJ > CO 5h •H U <-\ C N •H CO I— 1 l-^ N -H to i-H r-H Sh 0 QJ CO cx 0 ^•rf >.D 1—1 • QJ •H >, idi-l . X CO 4-1 4-1 r--^ E 4-1 CO >^ -^ E 4-J CO >> •iH x: 1-1 >^ c B :s >,-rA E C 5-1 4-1 •H •H E C 5-1 4-1 cn 5H rH b0 4-i •iH u U X 5-1 QJ QJ -H i-H X 5-1 QJ QJ •H en QJ -iH C •iH CO QJ m QJ CO QJ > > > C CO QJ > > > 0 > Sh 0 > s 4-1 0 > :s 4J QJ 0 -H ^3 S 4-) QJ 0 •iH Ph 0^ 1-H •H 4-1 cn 1 0 1 S-I 1 1 1 1 1 0 cn (U u 01 QJ d c 14-1 P u 5h cu •H E QJ C 5-1 X Cu QJ 1 0 5-1 CX QJ C i CX > 0 5-1 > 0 4J 0 > X QJ 0 > 0 0 1-H V-i 4J 5 CJ en 1-H 4-1 0 bO en Q) X ^4 ;:! QJ CO S-I ^) I-H Sh Mh QJ CO 0 X 0 •H QJ CL w 0 T) 0 W QJ ^ T3 U r-H P-i J-l (X^ CO Sh 1-h fx, 0) C > 4-1 cn en *. n C 0 4J QJ 0 0 > - 1 -a -H 4J •H C )-i C 4-J 4J - C E 4-1 0 QJ en i-H CO CO c: S QJ en CO •H cn 0) CO c QJ 0 E •iH c 4-1 fi 1=^ hi C C -rH E a 4-1 C 5-1 0 0 0 lU 0 0 13 QJ •H CO U QJ CO a •H cn •1-1 •H 5h QJ CO E x; Q) 4-1 4J CO 4-J 4-1 0 5h 4-1 e CJ 4J t— 1 0 0 CO QJ CO 0 0 bO CO b QJ CO < S C > E s CO 0 < T3 0 E nH -55- 1 -P 1 o 1 U 1 o C ::! O 4-1 C P o 4-1 •H JD 5-1 •H^ ■ 5-1 TJ fXr-l •T3 (Xi-t • - bO •H 4J QJ CO in •H 4-J •iH > CO 3 •H 4-1 •H > (U 5 (U bO g 5-4 g 5-1 S QJ bO g M g 5-1 c o u QJ ■H o •H CO 0 5-10) •H O •rH CO o kJ o u kJ a r~4 j: hJ O 5-1 ^J a.-i jc s CO CO CD i-l QJ QJ QJ CO C bO bO i-H QJ 1 • o c c e g CO CO C "-I >.-H CO O QJ CO 0) QJ QJ QJ x: 3nHx:4Jx;4-i.-Hx: 4J rH 4-1 1—1 ^ >> -H 4-1 CO O CO o bO w ^ bO CO Xi r-l 4-1 ^ i-H i-H QJ x: C ?-. CT3 4-1 C >^ CO 4-1 • 5h XI U-1 CO p 1—1 CO [5 c •H W N C 0 •H CO N C QJ O -H QJ T3 0) arH c O C •H QJ > C -H QJ > LM COi-H QJX; O^H OX) •H •^^ >^^-^ g •H •H >,i-l g •1-1 CO XI ex S ex c 4-1 CO x: CO QJ 4-1 CO ,C CO QJ 4-) ■U O g 5-1 C CO CO CO 4-J 4-) QJ bO O 4-) 4-1 QJ bO CJ a. cx QJ d •H 5-4 CO QJ 1-H CrH 5-1 CO QJ C 1-1 !-l CO QJ QJ 4-1 4-) CO QJ QJ 5-1 rH p •iH CO C •1—1 •H CO C •1—1 U X) CO CO g x: N CO ex CO QJ CO CO Xi CO QJ CO CO X) X C >^^H QJ 4-) •H C Q) O s^ •H g O X^-r^ g O W CO CO X! 5-1 O CO •H CO ex 1 1 >^ 1 c 4-1 4-1 bOi-i > •r^ cfi q CO c 1 C Xi •H CO 1 OJ -H 1 QJ -H 1 3 -1-1 -1-1 pH 4-1 g C > CO O > CO O bD4-l CO rH O QJ P 5-1 4-1 5-1 5-1 4J 5-1 QJ CO CO CO 3 5-1 CO C D- CO c a, 5-4 QJ O 5-1 X) >^ ^ -rH x: -H > a QJ O ^3 I-H CO g >. eg g >. >^ 5-1 > 5-1 l-H rH CO O 4-J > O CO - g x: 4J QJ .—1 4-1 •H 1-1 4-J •iH X) QJ CO QJ CO C bO p b0 4-> d b04-i X; QJ i-H rH 4J QJ C bO 60 C a bO bO C U 4-1 4-1 CO CO CO >^ CO CO 0) c O 3 QJ C O 3 •H CO x: QJ >^4-) CO x; Pi -H i-H X) P^; -H 1-1 T3 13 1-1 5 CO CO •H QJ O >^ >. 5-1 bOi-H 5-1 bOi-i 1 CO M-l O CO >^ C {X Mh o CO >.C a bO C 4-J O m (X rCl •H Ch O M-i cx rQ •iH a iH -H u CO •H 4J 13 CO •H 4-) 3 •H to u cu bOrC 4-1 CO C CO CO 4-J QJ box: 4-1 CO P CO CO ■u o ex C -H C CO C O QJ C -H C CO c O QJ CO C g QJ O •H c QJ bO-H >'0 QJ U -H C QJ bO^H >'T3 QJ -rH e c 4-) 5-1 O g QJ 4-1 O > c 4J Vi o > CO .'-S (u a c •H QJ •H CO CO O QJ a c •H QJ •H CD CO o 5-1 QJ 4-) t3 bO en :3 4J 5-1 CJ 1-4 ^ M-l bO CO Id 4J 5-1 O 1-1 JT MH to •H O cu CO o CO Cu 3 3 5-1 CO O to a 3 d 5-1 X; i-H QJ 3 c:--t O 1-1 -13 a QJ CO C 1-H a 1-H X) a. QJ CO fX 5-1 c CO I— 1 CJ QJ O QJ o > 4-1 CO 1-1 a QJ o QJ o > 4-) O g •H •rH ■u S CO CO 5-1 4-) 5-1 ex o •H S CO CO 5-1 4-1 U a o •H 2 •H X) o O •H X) X) 5-1 1 4-) CO 1—1 i-H CO bO C • cu CO QJ QJ CO CO O •iH Q) 4J •H in 4-1 > CO ' r— 1 . ■— s U -H U > •H -U CU 5-1 QJ O CO c QJ QJ 1-H CO o U JD CO (U rH CU g a. u o 1-1 g xi CO 4-1 CO •H QJ rH CU cn •H o 0) . •H a-H CO 4-1 cox; x; > 4D E C bO 5-4 -O 4-1 g h04-> o 5-1 bO O 5-1 d o o Q) CL, QJ O O QJ CO 3 o C c 5-1 CO H o a 5-1 ^-'l ^-\ CO O 5^ QJ cr g O •H p^ x; -56- M-l O c O •H 4-1 O . c CO o O O w OJ 01 > T3 •H 0) 4-) ^ OJ M C 1=1 H 4-1 QJ CO 4-1 •H .-n3 tt) C :i TJ C >^ Cfl 1— 1 0) C > o •H •H 4-1 U CO O i-< CO . >^ T) T3 :3 13 tJ 1 M-l !-i M-4 E 4J E iJ C -H O QJ 5-1 O CO O 4-1 CO O 4-1 CO CO '(3 E TJ T3 (U CO }-l T3 CO OJ M X! CO CU >^0JOC;>^C4-l>. >^ M-l Oi CO !-i C • M-l (U CO >-l C • bO,— 1 i-H 5-1 O TJ ID CO -U O E U ^ -HM-IOd i-l-H •H cu o 4-) >-i ,-1 M-l 4J % U Xi cn Q) :i U >> ^ CO CU d S-l >^ CUCCMOJ-'COCO-H CO CO O 4J tH 4-1 M-l Ph CO 4-1 -H 4-J M-l (X W 5-iQJO!i:co5:;>i^ 5-1 o o QJ 01 Q) o 0 C E QJ C CO OJ c •U >i CO J3 4-1 E g •H CO -rH ^ 5-1 CO 2 rj •r-l QJ JQ 5-1 • P >. E E rH Xl 5-1 -H 4-1 ID 5-1 4-1 CO •1-1 •H r-l QJ 13 CO CO 4-1 QJ CO X X CO E 4-1 CO QJ CO -P •H M-l CO CO E E d 0 5-1 •r^ CO Q o S s in •H M-l Cu bOT3 i-H X) OJ 4-J 1— 1 u CO CO CO o 4-) 4-1 •H a CO >^t3 C0T3 O QJ X) n-l 0) xs QJ 5-1 5-1 J2 4-1 4J QJ ^ -H ^1 ,c 5-1 E CO 0» O CO O o 0 E 4-1 c 4-1 CO CI, 4-1 a o CO o CO O QJ CO QJ cj) O z CJ D- 5-1 CJ 5-1 bO M c C •H •H M-4 o 5-1 5-4 •w •H o CO O CO T3 M-l ■u^ ■U ^ TJ 13 QJ ■rA •H o •r-l O QJ QJ 4-1 4-1 c o C O 5-1 4J 0 O C o 4-1 O 4-1 -H 5-1 QJ •H QJ CO E CO E CO ^J O r-l CO -H 4J cr P-r-l CO O CO O M-l O M-l QJ QJ 0 PQ cn Q s o Z O 5-4 p^ a 0 CO 1 q - 4-1 0) CO o & > .« 1 CO T3 -H 0 4-1 •rH c 5-1 QJ c -u 4-1 C C E 4J o 0) E .—1 CO CO c QJ CO CO •iH CO CO C QJ CO E -rH c 4J c C C C---I E 4-1 4-J d 5-1 CJ o o o o Ti QJ CO -H CO 5-1 QJ CO O •H CO •H -H 5-1 QJT3 E 4:: 0) 4J 4-J QJ 4J 4-1 O 5-1 E 0 4-1 1— 1 o o CO ^1 CO a o bO C b QJ CO < z c > 0 2 CO a < 0 CJ E --1 -57- 4-1 0) (U CO 5-1 C u C Q) QJ QJ 4-1 O CO -rH 4-1 O CO' •H CO 4-1 C W U-l 4-1 CO U-l 4J 1 O CO •H 0) 4-J CO QJ 4-1 CO CO n-( 4J 4-J 4-J > :>> O -H 4-J > >^ o •rH •H O CO CO d yt 4-1 c n 13 5-1 4-1 c 5-1 X) •H XI CO -H 'O ex X CO -I-l XI CX o:: 5h 4i c (U X C QJ .» 3 CO rH CX M d 1-H 4-1 CO . bO 13 ,-1 4-1 CO . -< ex C 5-1 >. t^ QJ t-H ex C 5-1 >^ >. g CO C C CU }-i =) o o (U X3 CO 5-1 3 O O QJ Xl CO •H x: 4J o bO O cx;3 o CJ bO O cx;3 o C U 4J CO c (U CO X w o C QJ CO X 4-J o •H ;3 o 13 >^ M S-l -H CO , CO CO 1 1 CO ^ CO CO 1 1 CO 15 CO .-1 o C g •- u E 4-1 CO rH • C e •- 5-1 g 4J CO ,-t • C CX CO O •H C Hi M-l QJ C 13 bO o •H c :3 M-l 0) C :3 bO o <-* C •iH O 4-1 4-1 OX O bOC •H o 4-1 4-1 OX o bO C XI CO o • 4-) CU O CO 3 •H QJ •H W QJ O CO :3 •H QJ •H •- C OJ •H XI •rH bO CO •H ^ (U QJ 4-1 5-1 4-1 •H bO CO •rH x QJ QJ 4-1 5-1 4-J g CO CO 4J QJ ^3 5-1 XI O O CO CO X3 5-1 X3 O O CO CO =) CO 4-1 TJ CO CO (U (i; C-i 5-4 QJ X) CO CO QJ c C--4 5-1 QJ g QJ >^r-l CJ CO r-( 4-1 (U O CO CO =! Q) > CO .—1 4-1 QJ o CO CO :3 QJ > •H I-H rH 13 QJ O E d C-H CXT3 5-1 O B c: C rH cxxi 5-1 C CO X P-MH o 5-1 CO o CO QJ CO O C CO O 5-1 CO o CO QJ CO o c CO •H X -rH O MH 2 O P. CO ^ 4-1 ^ CX I^X 2; O CX CO x 4-lX CX :3X s ;3 CO CX CO Ti •T3 QJ QJ C (U QJ r-l g O 4-) X) 4J XI X O 5-1 (U 5-J QJ CO CO C O N O N rH O Dh >^ CX >. •H CO •rH (U i-l QJ >-l cO QJ 4J S-i CO 5-4 CO > XI CO CO 3 g CO ^ CO X CO r-l 5h ^ o a QJ O O o 4-J Xl 4-1 XI C ,>.c O •rH O ■H . rH X O •U O 4-1 X5 4-1 o 4-1 Xl 4-1 •H •H •-•H CO QJ •H •-•rH CO QJ O X) 4J iUXlM-l bO bO CXIM-I bO bO QJ QJ CO O /--s 6 5^ 4-1 4-J CO 5-1 :3 B 5-1 4-1 •H C 4-1 CO 0 •H 5h Vh Xl O CU X) OJ ^ QJ o g cr-rH > o 4-1 P- C 3 o CU O c a o QJ O c c o ;3 o c •H CO ^4 CO •H QJ CO 5-1 CO •rH QJ 2 CO O C o c rH o •H 4-1 XI X) CX 5-1 i 4-1 CO rH CO bO . (U CO 0) QJ CO CO o •rH QJ 4-1 •rH yo 4-) > CO >-H /-v ■u •rH 5-1 > •H 4-1 cu 5-1 — 1 6X CO 4-1 CO •H QJ I-l eu en -H o QJ -I-l a.-'-i CO 4-1 COX X > ^ B C bO 5-1 XI 4-1 e bD4-i O 5-4 bO O 5h CO o O 'iAN : Absent 2 Convention, countries vnicn are planning uo narvest can go anead and narvest. MR. HOFFMANN: Sure. I ' n not saying leopardize tne negotiation of a conservation regime m order to acnieve tnis subsidiary, and admittedly suDsidiary, point that I've been talking about. I'm simply saying it is more of a proDlem tnan the attention that's been given to it in- dicates . MR. SCULLY: If I might just add a point in response, or at least in comment, it seems to me that there may be some confusion in your remarks between participation in the Antarctic Treaty and participation in the regime. It is our view that participation in the regime to conserve Antarctic living marine resources should not be limited to members, to consultative parties, or even to acceding parties to the Antarctic Treaty. It should be open to any state that is involved in the activities covered -- essentially, fishing activities . Therefore, to take the case of Brazil, and I'm quite sure I see the analogy; but it would be our hope that a regime could be designed to which Brazil would adhere to at such time as it began to undertake fishing activities in the area . To go back to the question of distributive elements, I think what bothers me to a certain extent is an assumption — an automatic assumption, which I think I hear, that krill will solve world food problems or will solve developing- country problems with regard to protein deficiencies. I'm not sure that's true. I'm not sure we're in a position to say that's true and to start negotiating on the basis of the fact that it is true. Krill may offer only a very high-cost protein for certain end uses. It may be that krill -- krills — will not compete favorably with other forms of fishmeal, for instance, or with various kinds of vegetable protein sources — soybeans or whatever — if it's being used as an animal feed or something of that sort. My reaction is that I don't know whether one should seek to establish or to rely upon Antarctic marine living resources as a major element in anybody's development strategy. We can't make that decision now. I would hope that the Convention will neither preclude that nor end up losing its conservation effectiveness by seeking incorpora- tion of elements that are premature. B-22 I would certainly hope that if it turns out that krill offers an important source of low-cost protein tnat protein would be applied to world food needs in some rational way. We can't maKe that assumption yet and we can't write that into a Convention yet. I would not like to see a Convention which in any way precluded that or in any way asserted any kind of exclusivity with regard to the question of resources being limited to x number of countries. Finally, let me close by saying that there are just a number of legal questions, juridical questions, raised by the point of assuming that resources in what we would consider to be high seas automatically have a common heritage kind of component . MS. MITCHELL: May I make one point? I don't think you need to proceed — or I don't think we're proceeding or sugges- ting that you proceed -- on the assumption that krill will have great potential for the developing countries. We just feel that it's necessary now to leave the door open to the use of krill later on for developing countries; and we feel that by not stating it in the Convention, however general a provision it may be, you are in fact precluding the option, because in- vestment decisions are being made and capabilities are being built up — plans are being laid — and it will be much harder later on to introduce any form of accommodation in the interests of the international community. THE CHAIRMAN: I don't understand your comments. Would you rather be more precise in an action that you desire that we undertake now? I would associate myself with Mr. Scully's views. I mentioned when we discussed this question before that I had no problem in leaving the Convention open to that kind of an arrangement, should it be decided upon later. But I personally had difficulty with the idea of bringing it forward now as a negotiating idea. I certainly do not pre- clude, and would personally favor, something equivalent to that part of Recommendation IX-1 which sets the stage for it and would allow it later. MS. MITCHELL: No, we're not suggesting that it be ne- gotiated now. We're suggesting that the possibility should be provided. THE CHAIRMAN: That causes me no problem. I just don't want to have to say that certain language is inserted specifically "for the purpose of". (Laughter) Any other comments? B-23 :S . DONNELLAK: Or. page 11 in the lES -- 10 and 11 -- are references to tne various conventions tnat do relate to cne Soutnern Ocean area or could. vjould it be possiDle — if I asK a foolisn question — somehow to include in tne regime tne means of urging ratification of particularly the pollution foolish type of conventions oy signatories to the conservation regime — somehow put into it some kind of wording that would put a bit of obligation on the part of the signatory states? Tnese conventions are not ratified, they are not in farce, and with increased activity in the area it might be important. THE CHAIRMAN: Yes. MR. SCULLY: I think in a hortatory sense, yes. You're not going to write a Convention that's going to have the affect, by its ratification, of bringing in force other conventions, but I think that certainly there will be pre- ambular language which would be appropriate for references to other international agreements, the desirability of their coming into force. In this area, however, we may not be in a position to cast too many stones. (Laughter) THE CHAIRMAN: We're not in the best shape that we could be on these agreements outselves. Other comments: Yes. MR. HOFFMANN: Maybe in closing -- I realize you might not want to answer this directly on the public record, but could you respond to Mr. Robinson's observation that you might make the negotiation process difficult by the openness of the EIS processes — what effect it might have on the negotiations, for example, in other countries that might lay their hands on the draft EIS? THE CHAIRMAN: I think it would be a help to the United States in the negotiations for others to see that the issues with which we're dealing have been widely discussed and that we have testified before the Congress on them. Indeed, I have already made the point in a diplomatic exchange that a particular U.S. view has in fact been publicly endorsed by the United States Executive Branch. I view this public meeting in the same way — that the attention that's given to the Antarctic Treaty and Antarctic issues is good. So I don't feel it will be a problem in the negotiations. MR. BARNES: Mr. Ambassador, I'm Chaplin Barnes of the National Audubon Society. I find myself in agreement with most of what's been said this afternoon; but I just wonder, as a final point: We spent a lot of time discussing the potential B-24 and exploitation of zhe Krill, and we're at the point of neaotiatina this regime Dasically oecause some countries are at a point of cmmercial development of krill. We're responding to exploitation pressures. I hope very much that tne U.S. delegation will write into the Treaty its basic purpose -- which is, in fact, to conserve the Antarctic ecosystem. This is the overriding purpose, and this is seen to be so in the document that you'd come back from Canberra with. I think it's very easy to lost sight of that in the details of the exploita- tion, limits, and so forth. i THE CHAIRMAN: That's certainly our intention. If there are no other comments, the meeting is adjourned (Whereupon, at 3:16 p.m. the meetng was adjourned) APPENDIX D Interim Report of the Second Special Antarctic Treaty Consultative Meeting at Canberra and Press Release of the Meeting D-1 ANTARCTIC TREATY P'^'^'JP TRAITE SUR LANTARCTIOUE SPECIAL CONSULTATIVE MEETING M#_Vi,>l3A KKLNION CONSLLTAIISL M'bUAlL 'E#J ZtOrOBOP OB ART^APKTHPCE ^^«^ TRATADO ANTARCTICO CntUnA.lbHOt KOHC.V.TbTATHBHOt CUbLLUAHllL ^ REUNION CONSULTIVA ESPtCL^L CANBERRA 16 .March 1978 INTERIM REPORT OF THE SECOND SPECIAL ANTARCTIC TREATY CONSULTATIVE MEETING 1. In accordance with the rrovisions of Article IX of the Antarctic Treaty, representatives of the Consultative Parties (Argentina, Australia, Belgium, Chile, France, Japan, New Zealand, Norway, Poland, the Republic of South Africa, the Union of Soviet Socialist Republics, the United Kingdom of Great Britain and Northern Ireland, and the United States of America) met in Canberra from 27 February to 16 March 1978 to consult together and consider measures which might be taken to further the principles and purposes of the Treaty and, where appropriate, make recommendations to their Governments. Specifically pursuant to Recoram.endation IX (2) of the Ninth Consultative Meeting, they met to elaborate a draft definitive regime for the Conservation of Antarctic Marine Living Resources. 2. The Meeting was formally opened by the Honourable Andrew Peacock, M.P., Australian Minister for Foreign Affairs. 3. Mr J.R. Rowland, Representative of Australia, was elected Chairman of the Special Consultative Meeting. Mr R.H. Wyndham of the Australian Department of Foreign Affairs was appointed Secretary-General and Mr R.L. Moncur of the Australian Department of Science was appointed Assistant Secretary-General o 4. The Opening Session was held in public. Opening statements were made by the Heads of Delegations (see Annex). D-2 5. The Meeting adopted the following Agenda : (1) Opening of Meeting (2) Election of Officers (3) Opening Statements (4) Adoption of Agenda (5) Elaboration of a draft definitive regime for the conservation of Antarctic marine living resources taking into account all the points listed in Recommendation IX-2 , Section III (6) Consideration of steps in connection with the work of the decisive meeting for the establishment of the definitive regime in Recommendation IX-2 , Section III (7) Adoption of Final Report (8) Closure of Meeting. 6. The Meeting considered in Plenary Session the first five items on the Agenda and appointed a Working Group, under the Chairmanship of the Representative of the United Kingdom, Sir Donald Logan, to consider the scope of the Convention to be elaborated, including the definition of the marine living resources of the area to which the Convention would apply. Elaboration of a Draft Definitive Regime 7. The Consultative Meeting discussed this item on the basis of eight draft Conventions and a number of working papers submitted by delegations and of the report and Recommendation IX-2 of the Ninth Antarctic Treaty Consultative Meeting held in London in September/October 1977. 8. After a full discussion of all aspects of the subject, the Meeting decided to ask the Chairman to prepare an informal text as a basis for its further work. D-3 9. As a result of detailed discussions of this text, the Representatives considered that it would provide a single text that would serve as a basis for further negotiation and decided to refer it to their Governments for study. They agreed that they would then continue their discussions and complete consideration of the Agenda at a resiomed session of this Second Special Consultative Meeting. In this connection they welcomed the invitation of the Government of Argentina to meet for this purpose in Buenos Aires in July, 1978. 10. The Representatives recalled the recommendation of the Ninth Consultative Meeting that a definitive regime should be concluded before the end of 1978 and felt that they had made very real progress to that end. D-4 -:'-nf -,0^; \,--wf^lf* -At^rNk^S^Sft ,\\'Vytiii^i,rf* ^yMi-i^fr^'^^'^9J\- <'^4'S-*.;.V^Vyp ■».:.■ ••\-:.' ''.'■if-- utiiitUr- kr ' II- 'i '■^- "'--"- ' •,^^i.«;.rJ.-.jf»#<-';i'.^5f:^V. Ne\A/S RELEASE •■S'...*- '^*'; .; - . -."it. - ^9" '-^;;'«''--'-.^< -. ■- . : '."f NQ D8 DATE 16 March 1978 1 f 0 r -' ^ < A ANTARCTIC MEETING The Special Antarctic Treaty Consultative Meeting was held in Canberra from 27- February to 16 March. Attached is the press statement released by the Meeting after the final session on 16 March. D-5 ANTARCTIC TREATY f^^^ TRAITE SLR LANTARCTIOUE SPECIAL CONSULTATIVE MEETING Mi -'I:'-"""^ RELMON rONSL LTA ri\ H slM ( lAM ZlOrOBOP OB AHTAPKTHKE ^^^^M TRATADO ANTARCTICO CnEUllA.lbHOfc KOHCy.lbTATIlBHOE t.UBtLUAHilt "* KEINION CON.-.LL1l\A ESPECIAL CANBERRA PRESS STATEMENT The first session of the Second Special Antarctic Treaty Consultative Meeting, which concluded in Canberra on 16 March, made very useful progress towards the elaboration of a draft definitive regime for Antarctic marine living resources. The main accomplishment of the Meeting was the production of an informal text which will be used as a basis for further negotiations at a resumed session of the meeting later this year. Drawing attention to the increasing interest being shown in the living resources found in Antarctic waters, the Meeting stressed the importance of such an agreement. The thirteen Consultative Parties - being those countries primarily interested in the Antarctic - had the main responsibility for the protection and preservation of the Antarctic and its living resources. In the 19 years since the Antarctic Treaty was signed, they had already taken a number of measures within the Treaty framework for the conservation and protection of Antarctic flora and fauna. They renewed their commitment to establish a definitive international regime for the conservation of Antarctic marine living resources before the end of 1978. D-6 A considerable number of proposals were examined at the Meeting, including the texts of a number of draft Conventions. These formed the basis of its work. The Chairman of the Meeting (Mr J.R. Rowland, Deputy Secretary of the Department of Foreign Affairs) tabled a single informal text reflecting the Chairman's assessment of the main elements in the various approaches suggested. This document was revised in the course of the Meeting to reflect trends which emerged as the discussions proceeded. The discussions revealed much common ground in the positions of the Consultative Parties and a common agreement that a Commission on the Conservation of Antarctic Marine Living Resources should be established. There was a wide measure of agreement on the functions of such a body. A common element in the positions of delegations was the need for the Convention to contain a number of agreed principles of conservation which should be applied by the Commission. It was also recognised that in carrying out its function the Commission would require the support of a Scientific Committee which would be a source of informed and objective advice to the Commission. There was also agreement that the Convention should reflect close links with the Antarctic Treaty. There would be a need to harmonise the activities of the Commission in the field of conservation of marine living resources with the responsibilities of the Antarctic Treaty Consultative Parties for the protection and preservation of the Antarctic environment. It would also be necessary to take account of other relevant international agreements and of international organisations such as FAO, whose co-operation would be important. Work still remains to be done on important aspects of the regime and in preparation for a decisive meeting to adopt a Convention. The meeting welcomed the D-7 invitation of the Government of Argentina for the session to continue its work at a res;amed session in Buenos Aires in July. The Australian Representative extended an invitation on behalf of the Australian Government to host at an appropriate stage a definitive meeting to adopt the Convention. APPENDIX E Role of Krill in the Antarctic Marine Ecosystem APPENDIX E ROLE OF KRILL IN THE ANTARCTIC MARINE ECOSYSTEM by Katherlne A. Green Box 1320^ El Paso, Texas 79912 Final Report to The Department of State Division of Ocean Affairs Contract 1722-7202^18 December 1977 C-1 TABLE OF CONTENTS page EXECUTIVE SUMMARY ill INTRODUCTION 1 QUALITATIVE ECOSYSTEM DESCRIPTION 2 Plants 2 Krlll 2 Other Zooplankton 4 Ice Invertebrates 4 Birds ^ Pish and Cephalopods 4 Seals 5 Whales 5 Benthos 12 Exchanges with Other Ecosystems 12 COMPARISON WITH OTHER MARINE ECOSYSTEMS .13 QUANTITATIVE ECOSYSTEM DESCRIPTION 15 Stock Estimates 15 Phytoplankton 15 Krill 15 Other Zooplankton l6 Fish 18 Squid 18 Birds 18 Crabeater seals 19 Whales 19 Predation Table 20 SUMMARY ECOSYSTEM DESCRIPTION 23 KRILL PRODUCTION 2l4 KRILL HARVEST QUESTIONS 25 Surplus Argument 25 Yield Estimates 25 MANAGEMENT AND CONSERVATION CONCERNS 2 7 REFERENCES 29 PERSONAL COrmUNI CAT IONS 3^ C-11 LIST OF TABLES page Table 1. Most Important fish species of the southern ocean 6 2. Seals in the Antarctic region 8 3. Whales of the Antarctic region 10 LIST OP FIGURES page Figure Conceptual model of the southern ocean marine ecosystem 3 Stock and krill consumption estimates Units 10" metric tons v/et weight 17 Annual total consumption. Units 10 metric tons wet weight 21 This study was funded by the Department of State through contract number 1722-720248. The ideas and conclusions expressed in this report are those of the author. They are not necessarily shared by the Department of State . C-111 EXECUTIVE SUMMARY The -Antarctic marine ecosystem is considered to be all waters south of the Antarctic Convergence with all the plants and animals living there. The ecosystem is defined biologically by the distribution of Euphausia superba (krill), some phytoplankton, penguins, and fish, and seals which are circumpolar but restricted south of the Conver- gence. Other species, most importantly whales but also some fish, birds, and perhaps cephalopods , migrate across the Convergence providing a link with other ecosystems. The Antarctic ecosystem is defined physically through the boundary of the Convergence, circumpolar upwelling and input of nutrients, transport by the Circumpolar Current, a uniform low temperature, fluctuating pack ice area, and the polar light regime. Most of the primary production is accomplished by phytoplankton, one celled plants. Algae in sea ice and some seaweeds also contribute. Krill are the dominant herbivore. Krill have an unusual ecological role since they are the main prey for fish, squid, Adelie penguins, fur, crabeater, and leopard seals, and baleen whales (fin, sei, blue, humpback, and minke whales). Other carnivores in the system not feeding on krill are toothed whales (sperm, killer, and some small whales) and elephant seals. Ross seals, Weddell seals, and some fish eat krill incidentally. Sei, minke, sperm and a few killer whales are now harvested in Antarctic waters. Several fish species are taken in commercial quantities. Squid, birds, and crabeater seals are not now harvested, but are potential commercial resources . The earliest estimates of production in Antarctic waters were made in coastal areas or at the pack ice edge, the most productive regions. Extrapolating those values to the whole southern ocean area produced unrealistlcally high values, giving the false impression that Antarctic waters were far more productive than other oceans. On the basis of later measurements in open ocean waters which are more representative of the area overall, estimates of total annual productivity have been revised downward. Although total annual plant production is not signifi- cantly greater in Antarctic waters than in other ocean areas, concentration in a short intense growing season and at the edge of moving pack ice makes production appear very large. Unusually short food chains allow a greater propor- tion of annual production to be transferred to carnivores, thus supporting very high standing stocks of mammals and birds . c-iv The ecological role of krill is unique. Krill have a longer life span than other marine zooplankton, requiring several years to reach sexual maturity. One species pre- dominantly, E. superb a, supports many different predators in five major groups, fish, squid, birds, seals, and whales. The dependence of many predators on krill as the main food resource implies that overexploitation of krill will be very damaging to most other populations within the ecosystem as well. If krill populations are reduced to the same ex- tent that whale populations have been, then the other poten- tially harvestable resources will also become scarce. Be- cause krill have a central place in the Antarctic marine food web, excessive harvesting would be dangerous to the entire system. The reliability of data on standing stocks of various Antarctic populations varies. The extremes of the food chain, mammals and plants, are the best known. Primary production averages about 25 gC/m^/year or 9500 x 10° metric tons wet weight of plants produced annually. This produc- tivity estimate is consistent with krill standing stock of 200 - 600 X 10° tons and an annual krill production of 350 - 400 X 10° tons. Krill population estimates are not based on direct sampling data, but instead are inferred from esti- mated predation on krill. Good quantitative estimates of krill abundance are not available, in spite of great scientific and commercial interest, but are needed. Direct estimates of fish and squid stocks are not available. Again population estimates are inferred from, predation estimates, and are not reliable figures. The estimates of 14 x 10^ tons for fish and 12 x 106 tons for squid in this report are speculative. Squid are probably underestimated . A standing stock of 0.27 x 10° tons of birds, mostly Adelie penguins, is a good estimate for Antarctic waters. Estimated populations of 5-5 x 10° tons of crabeater seals (25 million individuals) and 1.1 x 10° tons for all other seals combined are reliable. Estimated whale stocks derive from fishing data and are very reliable. Current stock estimates for whales are 4 x 106 tons fin, 1.4 x 10° tons minke, 0.8 x 10° tons blue, 0.7 x 10° tons sei, 0.08 x 10° tons humpback, and 1.16 x 10° tons sperm whales. Estimated annual predation on krill amounts to some 330 X 10 tons. Crabeater seals account for over 100 x 10° tons of the total. Squid may consume almost as much. Present baleen whales stocks take only about 43 x 10° tons. A commercial harvest of krill would compete with predators. Krill harvested will come from the 330 x 10° c-v tons estimated consumption, not from some hypothetical unused portion of the population. Potential harvest esti- mates range from 70 to 150 x 10" tons annually. A harvest at the higher end of that range would displace half of the predators, causing large changes In the Antarctic ecosystem. From an ecosystem perspective, a smaller sustainable yield, say 30 - 60 X 10° tons, seems more reasonable. It Is difficult to calculate potential krlll harvest from productivity of the stock Itself because of uncertain- ties In the life span and development rates of krlll. Whether the clrcumpolar krlll population consists of one or several distinct breeding stocks is also unknown. Management and conservation concerns on a krlll harvest from an ecosystem standpoint are a possible reduction in the rate of recovery of baleen whale stocks and limitation of eventual population totals, reduction in abundance of other potentially harvestable populations such as fish, squid and seals, and alteration of the trophic structure of the Antarctic marine ecosystem in the event of overexploitation because of the central position of krlll in the food web. A controlled harvest of krlll can provide some essential data on krlll population parameters and the response of the rest of the ecosystem to changing krlll abundance. In addition to usual catch and effort fishery data, nutrients, plant productivity, and populations of krlll predators should be monitored. C-1 INTRODUCTION Reviews of zoogeography of the southern ocean iden- tify the Antarctic Convergence as the northern boundary of the Antarctic Zone (Broch, 1961; Knox, 1968; Hedgoeth , 1969, 1970). Waters north of the Convergence are in the Subantarctic region. Some species of phytoplankton such as chaetoceros flexuosus Mangin or fragi lariops is sub linear is (van Heurck) Heiden are only found in the Antarctic zone (Hasle, 1968). The krill Euphausia superba are only found south of the Convergence (Marr, 1962). Antarctic seals and penguins probably do not migrate across the Convergence. Some fish and most whales do cross the Convergence to feed in the Antarctic every year. The Antarctic marine ecosys- tem is defined biologically by the species found in the Antarctic zone. The Antarctic marine ecosystem is also defined by physical oceanographic conditions which strongly affect its biology. The Antarctic Convergence is a physical fea- ture. Seasonal changes in available light and extent of ice cover are characteristic of the Antarctic zone. The Circumpolar Current serves to connect waters south of the Convergence into a single system through transport of nutrients and plankton. C-2 QUALITATIVE ECOSYSTEM DESCRIPTION A conceotual model diagram of the Antarctic marine ecosystem is presented in Ficrure 1. Boxes indicate eco- system components . Any organism found south of the Con- vergence can be placed into one cf the ccrxartrr.er.ts . Dissolved organic and inorganic nurrients and organic detritus are also included. These nonliving components are nuTrier." '::Is r.ecessary to comolete the nutrient cycles . In Figure I, bcx size is net related tc -ctal standing stocks of the compartments. Quantitative estimates cf standing stocks are given in Figure 3 . Arrows show material exchange ar-d rrophic (feeding) flows among ecosystem com.iDonents . The dark arrows repre- sent pathways with the largest annual total flows. They also indicate the most imDcrrant Dredator-prey relaticn- ships . Nurrienrs enter the sysrem through ur-.rellir.g . They are lost through A.ntarctic Bottom. Water f zr-.^zLzr. . through sinking to the sediments , and through harvest cr em.igra- tion of anim.al3 . Broken arrov/s represent the conrribution of dead organisms to organic detritus. 't::p lank ten prcvice SO to 90% of the primary pro- or carbon fixation, in Antarctic waters (El-Saved, sen, 1375). Phytoplankton are single celled planes. Dominant Antarctic species are diatoms (Holm- Hanser. , ee al. , 1977). Other plants in rhe ecosystem are algae growing in the bottom layer of one-year sea ice which may provide 10 to 15% of annual prim.ary production '?-r.t, 1953, 1968, 1971) and some seaweeds (BIOKiASS, 1977). -r^f^e Kr: Krill are euphausiid cruseaceans which are eaeen by baleen whales. They are classified as zooplankton or m.icronekton. The main species are Euphausi a superba , E. crustalloropkias , and Thysanoessa macrura (Nemotc , 1959). Frc.- ehe Antarctic Convergence south, the predominant krill species is Euphausia superba (Marr, 1952). Euphausia superba is the only krill species of commercial interest. Krill play a very important role in the dynamics of the Ajntarctic marine ecosystem. They are the main herbi- vore. '.-Jith zooplankton, rhe-" facilitate nuerier.e C-3 e 0) ^-> to CO o o (U c •H a c Id o o c 0) ■H O 0) 0 -O > -H (u q cA) P CJ X) 3 (0 0 4-J M -H C (C -M >. (T3 w pi q 13 QJ 'CO -rH 4-1 U £ c r— 1 (fl Q) U 13 0 -H ^-^ 0 > 0 (T3 ID J^ -P -H 3 CJ pj r^ ■P 0 fU f^ C W D C 0) < -H q W -rH [^ fl! 0 CU ro (fl C Q- 3 oj (X 0 >. 0) 15 q p en "^ P 60 0 r- -p 0 X3 iH E 0 q 1—1 ^0 C ^ -r-t ^ U 1 to m CO n Sh -P n 0) 0 0 4-) < (1) -t-' 1) fiJ .— 1 C 0) C -H QJ -p 3 •■ w •H zr W tD -H -H t-- 0 3 E QJ cn C9 -H cr on a fO 1 0 '+H X) U +-' (Ji -p X 0 A^ W X3 CO U 0 g 0 •H 0) ^ £ rH ■P S-, rfl 0 pi QJ PJ < rH 0 W D, (U bC 0 3 ^P pi a, m &, c S tU TD J- •• X3 0) X 6 3 C 0 0 Q) X n o •H , > •- S QJ 0 M 0 -H 1J rd -M 0 0 M r- rH -H >, Di •rH CQ ^ tl3 -iH 0) -rH m ■M 4:: 0 0) CU X3 PJ C CO P- ^ Mh 0) e bO >i 0 0 !h D. tn 0 0 f^ 0 i-H bC +-> 0 W 0) 0) X -H 0) 0 0 -H OJ a c Ph QJ Q) >, QJ 0 -H 4h 0 01 CJ CX 0) I4h f^ CL, 1+-,- fJ Sci 3 -H a. fn CJ ^ f-l Oh C/J c rd •" i-H ^ QJ M-H +-■ O c f^ bO .r 0 0 CX C x: s: x: -H < (D ■p pi pi c 1— 1 bCrH bC bC u E C C r-H c q Q) •H 6 0) -H QJ QJ ,C > r— 1 0 -H J^ 1— 1 rH -M bC w >, ^ 3 0 ■H C fXJ 0 e B G O rH D 3 5: r~- 0 w 0 u m 0 0 (TJ W 0 0 N (U 0) N PJ s: m 0 0 tT3 OJ 13 pj (U Ul U '• ^ u c rH 3 u •H U C (D bO 0 f-i 3 0 -H 0 4h 0 •H 2 -H tsl QJ U3 CO pi Q) 0 --H tfl e B C U D: C fn W «1 Xi 0 E X) -H ' ^ w w 0) nJ TD 0 C -'-H 3 c c 13 U c ffl .-H s: c a; nJ w P' m 13 QJ •H pj .c. 0 CD 0) 13 c 0 Oh bc q m •H U Zi -^^ , 0 0 fn QJ 0 f^ < •H P , Cfl C f^ •M 3 (D <-\ C •H fn x: QJ q fO •P 00 •- cr > OJ 0) X! 0 -H P 13 pi q -H -M m m 0 0 D e b3 C -P f^ 0 PJ 3 -X q f^ •H 0 f^ tfl 0 in fU D C 3 u q 0 S^ Q) 0 Q 0 < S CQ w S rH -H X3 00 < C/^ 0 Dh a, 6 •H «) 3 to 01 ■H ^ 3 3 OD "3 — , •H --H (0 —s 0 ■C ^ P" ^ q ' -c; fa cj Q' U) 0) (a 0 0 01 W to •H ■r^ Qj •r^ •-I w q ^ •H M --s tj 3 .^ C -(J 0 4h tfl -H S-l &1 q 'P PI -c; q ijh • Cfl Oj 0 0 --H 0) Qj -H (u q 0) -H 0 0 to m qj •— 1 S ^ e t) x: •-( ^ >H >H 0 3i -H 0 •H •u -H rg — V Q- 'C .^ 0 >-< -ij +j ■u 0) (C -N -O -U Ol -ri (1) 0 0 U) -u t) e q tn -u 3 ..^ 0 -Q -u T3 3 x: Is •— 1 0) ■u CO q 0 ti C q 0 « 0 ^ pi X) q 0 Oi 0 0 q CX 0 0 KC 0 -C 3 ^ •H 3 cy^ S 0 -H f:!: 0 to -H 0) (0 CO S M ~w 0 O^ v_ 5: T] -_ E: tJ,-- 0, q tr, ^ tH 1 C-7 C O o 0) rH ■p 0) CO r-\ ,^ •H /2 -H rd :3 > •p o (tJ .C CO bO 03 ;3 S-, +J (0 (fl (C O . TJ 4:: rH (/) c 0 ■H C m +-> C rO H (ti q; 6 -H 0 O ^ QJ Q) to ■P c < I a CJ •H to 3 (J •H •U) 10 O to to c (0 C to (1) o o C rH 0) (U O CO O to ■■H •'H +J <*H q »H x; o to -u to ^ o i c o nj n; -u ■P Q) ^ N '>-P O CO 3 C ^ 13) O O O C CO -H to I bO •^n-f O 0 6 rH en U 6 Q) 0 0 P" CD U W) to tjH rO +-I rH -p to C Q) X QJ Qj a, bOS >; -H 3 to 4:; to >, ■p u 0 Xi u >, to >.x rH r-\ a ^ C P- -H to ^ 6 to o S CO bo o C bO Ln -rl 0) I X C Pi O 3 000 ^ en o< H PJ to a CO -X) c c 0) to r-\ H a) CO bO 0) P" o to pi O CD C en •H pi O 0) C 03 lO CO TD PJ TJ C C 0) lO C rH CO c o to P" M PJ O lO o p> (0 c s lO < •H PJ bO ?H ex f^ 0) Q) o 4:: 0) P" Ci) o HJ PI O H to Q) q £X -H Q) 4) tn O -U tj »H to -N •U T3 q (U (U rH rH ^ XJ to to i-\ r-i •H •H to to > > to to to to P' p- to lO T) TJ X ^ CJ 0 pi PJ to to 0 0 0 0 c c LO I rH C to CO X) C 10 rH CO •H O < to H :3 (0 C H c P- lO o to O P- en to 3 > O tH Q) (0 CO rH O r-{ CX 03 e ^ 3 c O Q) ?^ C •H -H O P" P" O en to (0 p> Pi 'CO O >. C , (0 ^ H-i iH (0 iti C X) (0 < (0 X) e 4-' ^ rH o CO 0) O ^ H O ^ fH O CQ C o •H +-< •H +-> t/) •H O f^ :3 I +-> no fO 0) g w •'-H-' ,-\ •rH rH S •H m c ^ •H to -o 6 ti) X) tn O bC O 0) 4h fH fH o tu -p t/5 O tl) O H fn rO bO (L) O, C -P ■P O •H to >. to '-^ rH >i P' to Jh e tU T3 •H X f^ C Ph to to -H 2 c o •H Pi <0 ID TD 0) C fn to ft to • '- a) CO rH fn to >^^ s CO 00 U 0) (1) rH bO rH to -rH B :3 >, 6 ^ rH ,Q to X3 ip o o u a tl) o to U •H :3 O CO to r— I to +-> to to >i o pi TD -H -H Jh Mh rH to •P lO Oh C P' O bO fn OJ -H O rH CO 6 e -p o, 3 •-\ 6 o (U to r~- o £ O LH pi 00 P •H 3 5 '' to CO (U tu u o tu •H rH P" tp to o a; to o

nj (h ^ 0) a +j to c 0 HJ 0) 0 q 0) •H •o •-H « '•N U o 0 tj M tu .0 *H tS 13 fX 0 10 0 Q) CO t:i 0 ^ to bO I C C fn 3 O tu O -H C >i to P to o >, o - fn O c to rH TJ U lO +-> »> fn to to fn (X 0) P •^ tO to tu •-{ Xi p to to to tu fn tu to O to C to H fH ^ to bO 0) C >, tu >, p tu P bO to (0 tp to o tu t/1 o\o t>J f^ CU 1 tu o •-\ pi c to to H-" tu T3 13 /a C O to ^ X5 fH ^ to CJ to CU o •p o (0 10 0) to ttj X tJ tr, 3i M (h C t) 3 0 0^ In ■u 0 ■o Q. tl) D) H) -H tc M ^— to -^ XI CU to C ;3 X3 lO O 0 f^ ex X bC O CO rH -H rH to "P rH ^ ifl cx tu e (u e to U O CO f. >, O rH CO c to >,-H fn to rH to 6 rH pi tu rH •P to S r-i r-\ 4J •P 0 to to fn to tu to >^ •P 4-" I O tp :3 fn 0 ^ to to P o\o C CD fn < CO tu pi H-J to to pi tu tu tu r-\ =J Xa 0 {-, <^ 0 to c lO tu ^ fn to fn to c to U tu fn TD 0 bO rH X ID bO to c f^ 1 to fn <0 •H o-P -P tu to to 0 c o x: c bCK x: bC c r- (U bC •p C to C tu 1 fn •p •p bO •H f^ o to 4:: to rH C 0 ft Q) LO f^ C r-i •P •p ft ft •^ 0 tu y; pi 3 ip >. to •P CQ 0 ft bO CO O fH tu Pi -o fn C X (U c/:) lO r* c • to r< 0 U tu > fH c to CO H-J to. rH tO u rH PI tu to C C lO fn •p •'^ rP c to c 0) < 0 U fn tu tu X) X) c 3 ;3 HH It to pi X3 n} 0 tu 0 e to M 0) pi to •P ft O S 0 < ^ > C to bOpi tJ '-i 0 ■0 0 Q) x: to D, 0 ••H to 4J to CO to to 0 e 0 a; 0 u CO X3 c c to to tu CJ ^^ tO rH H-J rH CO •p 3 f^ fn -X U to bCTD r^ C 0 x: •P ft to X) 0 •H :3 r-t

-5 3: ' — C-9 -a c o o w O P. o ex o c •H -P 03 0) 0) S I o o Sh T) O C 3 O 1i CD bC C H f^ > Q) O t-l -(-' O I rO 0) U3 QJ f^ C •H CU P. cx o •u TD C 03 rH CO •H 0 ^ •H -p +J CD m H •H T? o c rH It W cx X 4:: (U -P 1 00 U rH 0) > c O-H •H ■M C 0) O C q o q S: -H C > 01 M Q) (U d) to ^ -H 1J 3 -Q U' M W 0) 3 C CO (t • ^ ti CO •H rH C Q) (fl O e CO CO o ■p to 0) O! 0) 4h O O C rC cp TJ iH C x: X^ ■P 3 CO O CO X3 O (t3 Pi C o .- e -H o o -p pi ip -p I o C CSJ 0) 3 I CO X3 LO bO O nj fO iH C •H 0) 0) CX t< rH X a c H c tC -H c H CU CO bO O ^ QJ >,-H C td nj J-< IT3 QJ e o x3 >> 3 0 C CO •H 4:: to -P CO e 3 T3 0 C .*% U^ (TJ Q) rH 0 C CO C 0 •H QJ bO C C fn 0 0 QJ •H > +-< It CO C It •H TD 0 rH hO QJ CJ) 3 U QJ CX 0 %4 4-, 0 QJ X) 0 CXCD to 3 M •0 —^ x: M a 11 Q) 13 Q) 0 ■-H to 0 M 4J 0) )h 0 N 3 u HJ "^H «i: &> ^— ^ 0) to -P •H (0 ip Xi o CO rH QJ XS X 3 O rH O O fn C •H •> TJ T3-H O 3 O CT' 4h CO I U bC QJ C > ■H O f^ QJ > O O CPi It QJ +J ^ >,-H rH O - 3 rH 3 O D< O rH X rH CO QJ o f^ -p QJ O c QJ bO cp f^ 0 QJ > ^ C ■p 0 u 0 0 c CJ •H to -P X) 0 c u It It rH +J CO c •H < (0 3 M 10 to ^ •c •H ■-H Q< -H <0 OJ m 0) 0 0 to 0 •H ■u Q< ^ 0 0 3 »H (h >*H •t -u C-10 en 3 en C 3 O S^ PQ XS C rO ,C O O c o •H bC 0) a •H ■P U ^ IX) +-> C < 0) I> ■H en iH O -^ W CO QJ < rH S fO O S OQ QJ rH C O •H -P -p X c o •H p XI •P P m •H Q o c C X5 < CO en O •H w 0) ■H O 0) a, 00 CD -P rHro ttJ O P' rH o rH :3 •rlOO > o •H ^ TJ C ■H 0) a, (fl p c 0) -H W P 0) -H J^ C CX-H QJ Pi CJ QJ P> O i^ 0) M (0 c ^ QJ 0 0) ^ •H > P bC (0 a) c S-i ••-■H >, 0 ^ U) •H o 4:: pi p pi 0 m c 5-1 f^ 0 tl bZ B p H C 6 J < o o Oj o 00 «^ o o " CM O O CO ^-- c; 10 •M u Ms QJ ^- -u 0. U) 0 3 q -H -^ Q) (0 OJ TJ to ■-H -^ Dl 15 ■tj ■C ■c: oq a s Q) P O QJ pi O a. 0) hO (C c Pj 0) O QJ ^ -H > P- K H ^ Q) ^— 4J Q< CO 0 3 C '-i -^ Q) 3 QJ It! 0 -H --H CO T) fO 3 ^ ID E S: TJ (13 0) Q) 0) > Q) QJ QJ CJ ^ X C P- QJ O bC ip P f-i rH OJ IJ TJ > -C QJ C e O •-DO > CD f-i CD IP O nj O P 13 ^' £ f^ O P b3 O 3 •H p O 6 CD ifi 00 en «^ O rH i_0 ''~N • lO o r- (Q 01 M to QJ ^— -u D. CO 0 ••H C ■-H -^ OJ to Q) fC QJ M f-H >^ t] 13 0 j:: CQ £1 s X} QJ P O QJ P O o p bC •H e o o r- cn OJ o o o 03 e 3 0) to 13 QJ ■U Qj -H OJ 13 "-H .-H tn tn u Q) 5: •X3 -a QJ - P pj CD C Q) 0 > CJ fn V-^ fU j: c 0 Sh -h (I3 bO >, QJ QJ t >, fn e QJ 0 •- x; -H >,P' p ^ 0 0 ip f-i P rH n) (t tfl p u x: c bC < •H QJ e bo QJ (B ^ CD t^ P CD QJ QJ > C rH rt) -H o o o O CM CM ^ — 13 — ^ 13 -u OJ (h 13 M QJ ^ 13 AJ ■u -c; a (0 S 0 0 C In QJ QJ 0 a; 13 •u c rH 3 ■H 13 CJ e CQ 1! C-11 c o u CO rH 0) ■H U Q) +J O D, >> rH H Iti ZS U CD 0) p ^ -M •- s:< rH 3 cti Uh c O CO •H C Cfl -H rC (D ro ■H O S U U (U O O J^ c 0 O c 0) x: bO -p fn 3 0) o > to C o (U o fn CO Vh ^ o M 1 Oj to -C 3 ■u — N t! 3 0) 0 '-H (0 0) ■TJ C ■^ •C r 0 CO •H QJ bO rH 0) n3 f-i e Mh 0 O 0) •H i-H +-' Xi J=) 0 h tJ f^ •H ■P 03 4:; •H ■P ■P rH C I Cu< (U X C OJ c 0 r •H o o o o to ^^N LO ro 00 J- ^ 1 0 ■u rfl pi t^ e 0 Ifl CO C ,c; pl c (1) •H CO C 3 O C X c 3 OJ m ■-H 0 to In •c 0 s w u 3 Q) C M •■H M 0 "H In ■*; 0 ^— to P" C .H (U iH TD 0) •H 3 CO 0) pj U O C >, rH >, X3 bO in o c Xi rH 3 C X) 3 QJ 0 e rH C 3 rH ^ CO (TJ c CO 6 3 m to C m Qj -H O H) pi to O C-12 (Laws, 1977)- Distribution of all species is circumpolar, with larger species and larger Individuals within species penetrating further south (BIOMASS, 1977). Benthos Benthic communities occur along the Antarctic conti- nental shelf and around islands. Energy sources for the benthic community come from seaweed production and from sinking of krill and other detritus from the water column. There is no commercial harvesting of Antarctic benthos at predeur. . Some seaweeds occur in harvestable quantities, particularly the brown algae Macrocystis pyrifera and DuTvillea antarctica and red algae such as the Gelidiales or Gigartinales . In the subantarctic , rock lobsters (janus sp.) are harvested. Several antarctic invertebrates (the scallop Adamussium colbecki , the clam Laternula elliptica , the gastropod Neobuccinum eatoni , and sea urchins Sterechinus sp.) are similar to species exploited elsewhere. However, small stocks combined with logistic difficulties indicate that commercial harvesting would not be practical. Exchanges with Other Ecosystems Carbon and nutrients are exported from the Antarctic ecosystem through migration of whales, some fish, and some birds. Nutrients are also lost through a presently small krill harvest, fish harvesting and whaling. A much larger krill harvest is anticipated in future years. In addition, other fish species, squid and possibly some seals constitute potential harvestable resources. The Antarctic marine ecosystem is well integrated. Component species and relationships between them have evolved together. There do not appear to be any empty niches. All the available resources are used within the system. It is very difficult for an outside species to in- vade a healthy ecosystem successfully. (Only systems under stress, such as the polluted Great Lakes, are susceptible to major faunal changes). Thus, importing a new species such as salmon to migrate across the Convergence, eat krill, and then return north to be harvested, is not expected to work on ecological grounds. In the case of salmon, there are also biological and physical oceanographic problems with establishing a population north of the Convergence. C-13 COMPARISON WITH OTHER MARINE ECOSYSTEMS There are several characteristics of the southern- ocean which are shared with all other marine systems: ( 1) Both organic and inorganic nutrients are dis- solved in the water column. This nutrient reserve is analogous to nutrients held in soil in a terrestrial ecosystem. (2) In land ecosystems, plant biomass is much greater than that of animals. This system exhibits an inverted pyramid of biomass; i.e.,' plant plankton standing stock at any given time is probably lower than animal plankton standing stock. However, the rapid turnover or replacement rate of the plant plankton allows the small standing crop to support the larger zooplankton biomass. As in other marine and terrestrial systems, the standing stock of other animal groups decreases with distance from plants in the food chain; i.e., standing stocks of primary carnivores are lower than standing stock of herbivores, etc . (3) Most of the primary production in the system is carried on by single celled plants. (4) Nutrients are recycled within the ecosystem, but energy is dissipated as it flows through the food chain. (5) Distributions of organisms are three dimensional in marine environments. Overlapping mapped distributions of two species does not necessarily indicate that they interact, since one may be a surface and one a bottom- dwelling organism. Krill are found at the surface in the southern ocean. (6) The biology of a marine ecosystem is closely tied to its physical environment. Light, ice cover, temperature and current movement are the most significant physical fac- tors in the southern ocean. (7) Productivity is higher in upwelling and coastal regions in the southern ocean as in other oceans. There are some special characteristics of the southern ocean which are not shared by other marine ecosystems and which deserve emphasis for understanding the dynamics of the Antarctic marine ecosystem. (1) Extreme seasonality in the light and ice regime, particularly below the Antarctic circle, dictate a very C-14 short but intense growing season. Physiological and behavioral mechanisms have evolved to spread the production pulse and to make food available throughout the year, even when the plants are not growing. (2) The southern ocean is productive, but not as much so as initially supposed. Early observations were made in coastal or ice edge areas during the summer. But on a per square meter per year basis, Antarctic waters are not much more productive than other oceans. (3) However, the short food chains in the southern ocean allow a greater proportion of fixed carbon to be transferred to carnivores such as fish, birds, and mammals. Since no group is more than three steps away from phyto- plankton in the food chain, the southern ocean supports an unusually dense population of carnivores. (4) Perhaps the most significant aspect of the sou- thern ocean ecosystem is the dependence of many predators on one prey species, krill. At first glance, ecological theory indicates that this situation should not occur; different predators tend to specialize on different prey species. However, the unusual dependence of several preda- to;."b on one prey is possible in the Antarctic because each pi'edator consumes a different segment or partition of the ki'ill population. Predators feed in different geographic locations, inside or outside the pack Ice zone, at different times of year, and take different size classes of krill. (5) The life span of krill is very long, compared to that of other marine zooplankton. In warmer waters, zoo- plankton may produce several generations per year. Krill, Ir contrast, require several years to reach sexual maturity. C-15 QUANTITATIVE ECOSYSTEM DESCRIPTION Stock Estimates Phytoplankton . Estimates of phytoplankton standing stock and production are derived from annual average pro- duction values for a small region extrapolated to the whole southern ocean. Production values are given in gC/m^/year, but most of the primary production occurs from November through February. The average ocean area south of the Convergence is 38 x lO^km^ (El-Sayed, in press). 2 An average production value of about 16 eC/m /year (Holm-Hansen, et al., 1977) gives 6080 x 10° metric tons wet weight produced. A conversion factor of 1 gram carbon to 10 grams wet weight for phytoplankton is used (Everson, in press). A production average of 25 gC/m^/year .Green, 1975) based on data for the Ross Sea produces a tctal pro- duction estimate of 9500 x 10^ tons. An estimate of 40 gC/m^/year (Currie, 1964) yields a production estimate of 15200 X 10^ tons. An average production estimate of 100 gC/m^/year was used by Ryther (1963). That value was a purely speculative estimate based on the concept that Antarctic waters are significantly more productive than those of other oceans. It has been shown by field data collection to be unrealist ically high. All of the values given here measure net carbon fixa- tion rates. Losses to respiration and- losses to leakage of organic matter have already been accounted for. The num- bers presented here are fixed carbon which becomes avail- able to herbivores. Krill. In spite of the commercial and scientific interest in krill, accurate estimates of krill abundance are not available. The lack of information is partly due to sampling techniques. Normal zooplankton collecting nets do not caoture krill effectively since krill are stronger swimmers than most zooplankton. High speed trawls, such as those used in the fishing of krill, are more appropriate for quantitative sampling. They have not been used over a wide enough area to provide estimates. The swarming habit of krill makes their distribution extremely patchy. It is difficult to extrapolate a catch inside of one swarm to the southern ocean as a whole. Participants at the SCAR Conference on Living Resources of the Southern Ocean in Woods Hole. Massachusetts in 1976 discussed total krill biomass in the 200-600 x 10^ tons range. On the basis of 75 x 10^ tons dry weight C-16 (Gulland, 1970) for krill, a wet weight biomass of 225 x 10 tons can be calculated. Working on a production to biomass argument, with a production to biomass ratio of 1.8:1 (Allen, 1971) and a production estimate of 330 x 10^ tons as derived from the chart in Figure 3, a biomass estimate of 183 x 10^ tons is calculated. High range of krill biomass estimates is represented by 930 to 1950 x 10^ tons (Makarov and Shev- kov, 1972). Krill biomass estimates represent a summer or maxi- mum standing stock. Annual average values are lower. For the remainder of the paper, discussions and calculations use the value of 25 0 x 10^ tons for an annual average and 600 X 10^ tons for a summer maximum biomass. Figure 2 lists estimates of population sizes for the compartments of the Antarctic marine ecosystem as in Figure 1. Phytoplankton production values are reliable. Exten- sive data collection over the whole southern ocean has contributed to the estimates. Recent estimates center around the range of 25 to 35 gC/m /year, similar to produc- tion values in other oceanic waters. However, in the Ant- arctic, production is concentrated in the summer season rather than spread throughout the year. Krill estimates are not reliable. They are based on speculations, not quantitative data. However, an estimated standing stock range of 250 to 600 x 10" tons and an annual production rate of 300 x 10" tons are compatible with phytoplankton production estimates. A primary production estimate of 25 gC/m^/year yields 9500 x 10° tons available to herbivores each year. If half of that is consumed by krill with only a 10% assimilation efficiency, an annual krill production of 450 x 10^ tons can be supported. Other Zooplankton Because of the difficulties of sampling zooplankton in patches and of taking net tows in pack ice , reliable estimates of non-krill zooplankton are not available. It is usually assumed as in Gulland (1970) that the biomass of other zooplankton is the same as that of krill. This estimate is a very soft number and not reliable. However, the lack of reliability is not significant because of low utilization of non-krill zooplankton in the food chain. An indication of relative abundance of krill and other zooplankton would be useful. A specific population esti- mate, however, is much more important for krill than it is for other zooplankton. C-17 STOCK ESTIMATED AVERAGE POp-JLATION SIZE X BODY WEIGHT EATEN^ ANNUAL FOOD CONSUMPTION TOTAL KRILL OTHER Phytoplank- ton Production 6080 (16 gC/m'^/yr: Holm-Hansen, et al 1975) 9 5 00 (25 gc/m2/yr: Green 1975) 15200 (40 gC/m^/yr; Currie 1964) assumes area 34 x 10^ km^ and 1 gC ; 10 g wet weight Krill 200 - 600 (Woods Hole conference) 225 (75 dry wt.; GuUand 1970) 183 (P-B 1.8:1: Allen 1971, and 330 as P estimate, chart) 930-1350 (Makarov and Shevkov, 1972) calculations used 250 ave. and 600 max Other zoopl same as krill (GuUand 1970) Fish 14 (Green 1977. based on preda- tion estimates) 6.5 Everson 1970 91 64 27 Squid 12.4 (Green 1977, based on preda- tion estimates) 11 Hurley 1976 136 100 36 Birds 0.2 7 (Prevost.in press) 67 18 14.' 3.6 Crabeater 3 (Laws 1977) U-6 (Siniff and Hofman, pers . com. calculations use 5.5 23.45 Oritslanc 1977 111 106 5 Other seals Ross 0.38 (Laws 1977) fur 0.015 (Laws 1977) elephant 0.3 (Laws 1977) Weddell 0.25 (Hofman, p.c.) leopaid 0.2 (Hofman, p.c.) total 0.8 23.45 18.: 3.1 15.6 Baleen whales blue 0.8 (Laws 1977) fin 4 humpback 0.08 sei 0.7 minke 1.4 4.2 (Lockyer 1976) 14.6 3.5 16.9 0.3 3.0 20.4 3.4 16.4 0.3 2.9 19.8 0.1 0.5 0 0.1 0.6 Toothed whales sperm 1 .16 (Laws 1977) killer 0.05 (rough guess) 4.2 25 4.9 1.25 0 0 4.9 1.2^ Fig. 2. Stock and krill consumption estimates, Units 10^ metric tons wet weight . C-18 Fish . The fish abundance estimate in Figure 2 is an indirect calculation. The amount of fish consumed by fish predators, as in Figure 3, is 14- x 10° tons per year. Derivation of this value will be discussed later. Assuming conservatively that annual production is equal to standing stock for fish, biomass is 14 x 10^ tons. If annual pro- duction is less than the average standing stock, then the biomass estimate should be revised upward. From a study on fish in the South Orkney Islands (Everson, 1970), an annual consumption rate of 6.5 times body weight is derived. A fish stock of 14 million tons then consumes 91 x 10" tons of food per year, about two- thirds krill and the remainder other organisms. Values for fish standing stock and krill consumption are educated guesses, and are most likely to be underestimates. Squid . The biomass estimate in Figure 2 is based on predation on squid, another indirect estimate. Previous estimates of Antarctic squid biomass have been based on the population size required to support predation by sperm whales, a major squid predator. However, from Figure 3, seals, birds, and possibly fish together consume at least twice as much squid as do sperm whales. Therefore, squid population estimates have been revised upward (Green, 1977). Estimated yearly predation on squid in the southern ocean amounts to about 19 x 10^ tons. On the assumption that squid populations turn over faster than fish do and that annual production is one and a half times standing stock, a squid standing stock of 12.4 x 10^ tons was derived. This estimate is speculative S.nd not reliable. It is higher than previous estimates, but may still be conservative. An annual consumption rate of eleven times average body weight is extrapolated from laboratory studies (Van Heukelem, 1973; Hurley, 1976). The total consumption by a stock of 12.4 x 10^ tons of squid would be 136 x 10^ tons of food per year. Squid may consume 100 x 10 tons of krill annually (D. Tranter, Woods Hole Conference). Fish and squid estimates are the least reliable num- bers in Figure 2 since they are not based on any direct data. One concern expressed at the Woods Hole Conference was the need for improved data on fish and cephalopod stocks. This information is essential, because fish and squid feed extensively on krill and they are potentially harvestable resources. Birds. From an extensive review of population and physiological data available for Antarctic birds (Prevost, in press), a standing stock of 0.27 x 10^ tons for waters C-19 south C'f the Convergence was estimated. The bulk of this biomass is Adelie penguins which subsist primarily on krill. Antarctic birds consume 67 times their body weight per year, .for a total prey consumption estimate of 18 x 10 tons. Proportions of krill and other species in bird diets were given for the Antarctic and subantarctic regions combined. Correcting these ratios to allow for the dominance of Adelie penguins south of the Convergence, birds consume 14 . 4 x 10" tons of krill annually. Because birds are easy to census, since they breed in colonies and in accessible land areas, the bird biomass and consumption data are very reliable. Birds have a relatively large impact on the ecosystem in proportion to their standing stock because of their high consumption rates. However, compared to seals and other species as krill consumers, birds have a relatively small impact. However, penguins may prove to be useful indicators of krill abundance (Green, 1975). Data on penguin popula- tion size and breeding success can provide an important key to the condition of krill standing stocks. Birds are much easier to sample than are zooplankton. Crabeater seals. Crabeater seals are by far the most abundant of Antarctic seals. Population estimates vary from 15 million individuals (Laws, 1977) to between 20 and 30 million individuals (Hofman, conversations at Woods Hole Conference) to estimates of 30-70 million individuals (Erickson, et al . , 1971). For the rest of the paper, cal- culations use 25 million indiiriduals or 5.5 x 10^ tons of crabeater seals. Using an annual consumption rate of 23.45 times ave- rage body weight (Oritsland, 1977), total consumption by crabeater seals is 111 x 10^ tons of food. Since their diet is 94% krill (Laws, 1977), they eat 106 x 10^ tons of krill annually. Biomass estimates for other seals are from Laws (1977) and Hofman (discussions at Woods Hole Conference). Total biomass estimated for other seals is 0.8 x 10 tons (com- Dared to a total of 5.5 x 10^ tons for crabeater seals alone). Other seals consume a total of 18.7 x 10° tons per year of which 3.1 x 10 are krill (after Laws, 1977). Whales . Baleen whales found south of the Convergence are blue fin, humpback, sei , and minke whales. Fin whales are the '?:^st abundant with a total estimated biomass of 4 X 10^ tc:^. Minke vjhales , although less abundant, consume more bec-^.:se they feed for a longer period of time in the C-20 Antarctic than fin whales. Present annual consumption of krill by whales is approximately 42 x 10^ million tons (Laws, 1977), less than half of the total consumption by seals and also less than consumption by squid. Sperm and killer whales are present in the Antarctic ecosystem, but do not consume krill directly. They eat fish and squid and are one step removed from krill in the food chain. Data on whales are based on fishery statistics and are very good estimates of population size and consumption rates. Seal information is almost reliable as that of whales. Both aerial and shipboard techniques have been used to estimate seal populations. In Figure 2, phytoplankton production and bird, seal and whale standing stocks are the most reliable numbers. Krill and other zooplankton estimates are considerably less reliable. Fish and squid estimates are the softest numbers in the table, essentially educated guesses. Predation Table Figure 3 shows estimates of the total biomass flowing along the arrows of the conceptual model of Figure 1. Values are in units of 10° tons and represent annual totals for the area south of the Convergence. Flows are located with the donor compartment as column heading and the recei- ver compartment as row heading. Flow from krill to penguins, which represents consumption of krill by penguins, is in the krill column and penguin row of the matrix and is 14.4 x 10" tons per year. Ice algae production is about 5% of phytoplankton pro- duction (Green, 1977). Ice invertebrates feeding on ice algae provide a mechanism for putting that carbon into the water column. Zooplankton and fish larvae eat ice algae. For the higher trophic levels , the matrix has been derived from the predator perspective. Estimates of predation on krill by fish, squid, pen- guins, seals, and baleen whales were documented in the dis- cussion of Figure 2. Consumption of other zooplankton, fish, squid, and birds are from Green (1977), Laws (1977) and Prevost (in press). Bird and mammal consumption rates are based on exten- sive data. Fish and squid consumption rates are specula- tive . C-21 TO FROM 0) (U 0) hC O r-H M < 1 -p S-. 0) (D > U C h- 1 1— 1 c 1 o O -P -P X > c X 11 rH r-H •P JnrP 0) 0-. x: o -p o OtS] X. •H •H 3 cr oo -a • p CQ Ice Algae Ice Inverts . 90 Phyto- plankton Krill 75 2500 Other Zoopl . 75 U 2000 Fish 10 64 11 6 Squid 100 34 2 Penguins 14 . 4 1.8 1.8 Crabeater Seals 106 3 2 Other Seals 3.1 1 7 4.7 0.5 Baleen Whales 42.8 Toothed Whales 0.5 4.6 0.3 Figure 3. Annual total consumption. Units 10 metric tons net weight. C-22 Figure 3 represents the best present estimates of con- sumption within the Antarctic ecosystem. The matrix indi- cates that squid and crabeater seals are the most important consumers of krill. Baleen whales consume only about a fifth of the total taken by squid and crabeaters . C-23 SUMMARY ECOSYSTEM DESCRIPTION The term krill refers to several species of zoo- plankton in the Antarctic ecosystem. The dominant species, and the only one of commercial interest, is Euphausia superba . Krill is sometimes used as a synonym for this species. E. superba is rarely found north of the Antarctic Convergence, but is abundant immediately south. Some fish, bird, and seal species exhibit the same distribution pattern, circumpolar south of the Convergence. All waters south of the Antarctic Convergence belong in the Antarctic ecosystem. The Antarctic marine food web is centered around krill. The Antarctic ecosystem is unusual ecologically since one species, krill, provides the major food resource for fish, squid, birds, crabeater seals, and five baleen whales. The timing of whale migrations, the fluctuation of the pack ice, and the relationship of seal distribution to pack ice type provide for some partitioning of the krill resource. The partition taken by fish and cephalopods is not known, although both groups are thought to be significant preda- tors on krill. Krill play a key role in the nutrient dynamics of the Antarctic marine ecosystem along with other zooplankton and phytoplankton. They facilitate nutrient regeneration in the euphotic zone during the growing season. Harvesting of krill, fish, and whales removes nutrients from the Antarctic ecosystem. Nutrients are brought into the system by exten- sive upwelling. The biology of the Antarctic ecosystem is closely regulated by the extreme seasonal variations in availability of light and in areas of sea ice. The circumpolar current unifies the system through transport of nutrients and plankton. Krill have a circumpolar distribution as do most birds and mammals. The distribution of fish and squid species is less well known. Whales and at least one fish species migrate across the Convergence. Squid are probably most abundant near the Convergence, and may cross it. C-24 KRILL PRODUCTION Total predation on krill by fish, squid, birds and mammals amounts to approximately 330 x 10" tons per year. This value is an estimate of production by the krill popu- lation . The ratio of annual production to average biomass is an estimate of turnover rate for the krill population. Pro- duction estimated by predation is roughly equal to the ave- rage standing stock and about half of the peak annual bio^ mass. Turnover of 50 to 100% of the population in a year is slow for marine zooplankton. However, Antarctic krill are long-lived in comparison with other zooplankton, so this figure is not so surprising. The questions of krill life span and age of sexual maturity are not yet resolved. Data on size distribution can be interpreted in several ways (Fraser, 19 37 •, Bargmann , 1945; Ivanov, 1970). Krill do not breed until the beginning of their third or fourth year, lengths of 28 to 45 mm, and may breed one or two seasons (McVJhinnie, letter to K. Green, 18 October 1977). Life span questions affect estimates of krill replace- ment rates and consequently estimates of sustainable yield. With present uncertainty on development rates, the propor- tion of the population that is capable of reproducing in any year cannot be determined. C-25 KRILL HARVEST QUESTIONS Surplus Argument Mention of krill as a harvestable resource is invaria- bly connected with an observation on the decline of whales in this century. The argument is essentially that an amount of krill equivalent to the difference between present con- sumption and peak consumption by whales can be harvested. Assumptions associated with this argument are that man can exactly replace whales as a predator in the Antarctic eco- system and that consequently harvesting will have no impact on the rest of the system. This argument is no longer con- sidered valid by scientists, but remains a part of the pub- lic attitude on Antarctic resource potential. There are several very strong arguments against the assumption that krill harvesting will not affect the Antarc- tic marine ecosystem. First, harvesting can not exactly replace whales as a predator. The fraction of the krill population which the whales consumed, with a specific cir- cumpolar distribution, time of year, size of krill removed, and relationship to pack ice, will not be duplicated by commercial harvesting. Second, there is evidence that the ecosystem has already adjusted partially to the reduced whale population. Penguins and some seal populations appear to have increased to take advantage of more available krill (Conroy, 1975; Laws , 19 7 7b) . A large commercial krill harvest will compete with krill predators . V/ith the change in abundance of krill available to the ecosystem, there will be changes in preda- tor populations. It is suggested that populations will return to 1900 levels -- i.e., to what they were when whale populations were considerably higher (J. A. Gulland letter to K. Green, September 1977). Population levels for fish, cephalopods , birds and seals around 1900 are not known, however. Yield Estimates The estimate of total predation on krill from Figure 3 is approximately 330 x 10 tons. Any commercial harvest will be taken from those 330 x 10 tons. Harvesting will displace some predators. Literature estimates of sustainable krill harvest range from 70 to 150 x 10^ tons per year (Moiseev,- 1970; C-26 lyfeLCklntosh, 197O; Everson, in press). Public expectations are even higher (e.g., 200 x 10° tons, Gwynne, 1977). Changing the perspective from only krlll and whales to the whole ecosystem, estimated harvesting levels appear to be too large. A harvest of 150 x 10° tons would displace half of the current predators, producing dramatic changes In the ecosystem. The estimate of 330 x 10° tons for krlll production Is a sum over the whole southern ocean area. A local harvest, say 10 x 10° tons In the Scotia Sea, would probably cause observable changes In predator populations there. My present educated guess for a sustainable yield, based on possible replacement rates and ecosystem Interactions, Is 30 to 60 X 10° tons per year for the southern ocean, with the larger value less likely to be sustainable over a long time. To try to maintain balance within the food web, preda- tors could be harvested In proportion to prey (krlll) harvested. Sustainable harvests of seals, birds, fish and cephalopods cannot be estimated well at present because of lack of Information on replacement rates within each of those stocks. As a rule of thumb for general consideration only, a total predator blomass of about 10^ of the total krlll harvest could be taken. Thus, 30 x 10° tons of krlll harvested would imply a total of 3 x 10° tons for seals, birds, fish and cephalopods. About half of that total would be crabeater seals. A concomitant harvest would necessitate management of all the resources, including krlll, by one agency. To meet the objective of maintaining a balanced ecosystem, the geographic locations of all the harvests would have to be coordinated as closely as the total removed. Predators should be removed from essentially the same location as prey. Population dynamics within each species must also be taken into account. C-27 MANAGEMENT AND CONSERVATION CONCERNS Management concerns from an Antarctic ecosystem per- spective in the area of environmental sensitivity to a krill harvest are: (1) recovery rates of baleen whale populations. Baleen whales are not a significant predator on krill at present. Even when whale populations were greatest, baleen whales probably consumed only about half of the krill avail- able to predators (Green, 1975). Recovery rates of baleen whale populations depend on the age ^t which whales reach sexual maturity as well as on total population numbers. Whales now become sexually mature sooner than they did at the turn of the century (Gambell, 1976). This promotes a rapid recovery of the population and may be due to good physiological condition from abundant food supply. A reduc- tion in krill might cause a rise in the age of first repro- duction and, consequently, slower recovery of the baleen whale populations. In addition, a harvest will be competi- tion for food, limiting the potential sustainable baleen whale population size. (2) abundance of potentially harvestable fish and squid. Fish and squid are potentially harvestable resources. They are probably easier to market than are krill even though they may be more difficult to catch. Some commercial fish- ing is already occurring in Antarctic waters. A substantial krill harvest could interfere with existing and potential fisheries by reducing prey availability and consequently overall population size, (3) populations of penguins and seals. A large krill harvest acting as a competitor for penguins and seals could easily result in a reduction in penguin and seal populations. The most affected would be crabeater seals, a potentially harvestable resource. (4) trophic structure of the Antarctic marine ecosys- tem. The harvest of krill is not analogous to the harvest of whales. A whale is a top carnivore or an end of a food chain. Some baleen whale populations were reduced to less than a fifth of their highest levels. While there have been adjustments within the Antarctic ecosystem to changes in whale populations, the basic structure of the system remains the same. If krill populations were reduced com- parably, the entire character of the Antarctic ecosystem would be changed. Some additional information is necessary for the conservation and manapeTnp>nt n-F krill stocks. Breedinor C-28 stocks must be identified. Data is needed on the abundance, distribution, age structure, and reproductive physiology of krill. Limited commercial harvesting can provide some of the data necessary for calculating sustainable yields. In the event of any krill harvest, data should be collected, reported and analyzed: location, time, search time to locate a swarm, gear used, swarm size, shape and depth, size composition of the catch, and breeding condition. Monitoring the rest of the system for a response to the krill harvest is also essential. Nutrient levels and primary production rates in the vicinity of krill harvesting should be measured. If possible, fish and cephalopod popu- lations should be evaluated. Birds, particularly penguin populations, should be monitored closely as potential indicators of changes in krill abundance. Seal and whale data are already being collected under existing conventions. A controlled krill harvest can be treated as an exper- imental manipulation of the Antarctic marine ecosystem. With careful observations, data pertinent to the ecology of the system as well as to management of the fishery can be obtained. C-29 REFERENCES Allen, K. R. 1971. Relation between production and biomass. J. Fish, Res. Bd. Canada, 2Q_: 1573-1581. Andriashev, A, P. 1968. The problem of the life community associated with the Antarctic fast ice, In: Symposium on Antarctic Oceanography, pp. 14-155. R, I. Currie, Editor. Scott Polar Research Institute Cambridge, England. Bargmann, H. E. 1945. The development and life history of adolescent and adult krill, Euphaus ia superba . Discovery Reports, 23: 103-178. BIOMASS 1977. Biological Investigations of Marine Antarctic Systems and Stocks. Volume 1: Research Pro- posals. Scientific Committee on Antarctic Research. SCOR Working Group 54. Broch, H. 1971. Benthonic problems in Antarctic and Arctic waters. Sci. Res. Norwegian Ant. Expeds . , 38: 1-22. Bunt, J. S. 1963. Diatoms on Antarctic sea-ice as agents of pri- mary production. Nature, 199: 1255-1257. 1968. Microalgae of the Antarctic pack ice zone, ln_ Symposium on Antarctic Oceanography, pp. 198- 218. R. I. Currie, Editor. Scott Polar Research Institute, Cambridge. 1971. Microbial productivity in polar regions. Symp. Soc. Gen. Microbiol., 2]^: 333-354. Conroy, J. W. H. 1975. Recent increases in penguin populations in Antarctica and the subantarctic , Iri: The Biology of Penguins, B. Stonehouse, Editor, pp. 321-336. University Park Press, Baltimore, C-30 Currie , R. I. 1964. Environmental features in the ecology of Antarc- tic seas, In: Biologie Antarctique, pp. 87-94. R. Carrick, M. Holdgate , and J. Prevost, Editors. Hermann, Paris. El-Sayed, S. Z. 1970. On the productivity of the Southern Ocean, In: Antarctic Ecology, Vol. 1, pp. 119-135. M. W. Holdgate, Editor. Academic Press, New York. In press. Primary" productivity and estimates of poten- tial yields of the Southern Ocean, In: Polar Research: To the Present and the Future, M. A. McWhinnie, Editor. AAAS . Erickson, A. W. and R. J. Hofman 1974. Antarctic seals. Antarctic Map Folio Series, Folio 18: 4-13. American Geographical Society, New York. Erickson, A. W. , D. B. Siniff, D. R. Cline and R. J. Hofman 1971. Distributional ecology of Antarctic seals. In: Symposium on Antarctic Ice and Water Masses, Tokyo, September 1970, pp. 55-95. G. Deacon, ed. Scientific Committee on Antarctic Research, Cambridge . Everson, I. 1970. The population dynamics and energy budget of Nortothenia neglecta Nybelin at Signy Island, South Orkney Islands. Brit. Antarctic Surv. Bull. , 23: 25-50. In press. Resource review - krill. In: BIOMASS,. Vol. 2: Background Papers, S. Z. El-Sayed, Ed. Scientific Committee on Antarctic Research. Eraser, F. C. 1937. On the development and distribution of the young stages of krill (Euphausia superba) . Discovery Reports, 14: 1-92. Gambell, R. 1976. A note on the changes observed in the pregnancy rate and age at sexual maturity of some baleen whales in the Antarctic. ACMRR/MM/SC/37 . Food and Agriculture Organization of the UN. C-31 Gordon, A. L. 1971. Oceanography of Antarctic waters, Antarctic Res. Ser.", 1^: 169-203. Green, K. A. 1975. Simulation of the pelagic ecosystem of the Ross Sea, Antarctica: A time varying compart- mental model.. Ph.D. Dissertation, Texas ASM University, 157 pp. 1977. Antarctic marine ecosystem modeling. Marine Mammal Commission Report No. MMC-76/03. National Technical Information Service PB 270 375, 111 pp. Gulland, J. A. 1970. The development of the resources of the Antarc- tic seas, Iri: Antarctic Ecology, Vol. 1, pp. 217-223. M. W. Holdgate, Editor. Academic Press, New York. Gwynne , P. 1977. Get ready Antarctica - here comes the boom. International Wildlife 7(6): 4-10. Hasle, G. 1968. Marine diatoms. Antarctic Map Folio Series, Folio 10: 6-8. American Geographical Society, New York. Hedgpeth, J. W. 1959. Introduction to Antarctic zoogeography, in dis- tribution of selected groups of marine inverte- brates in waters south of 35°S latitude. Antarctic Map Folio Series, Folio 11: 1-9. American Geographic Society, New York. 1970. Marine biogeography of the Antarctic regions, In Antarctic Ecology, M. W. Holdgate, ed.. Vol. 1, pp. 97-104. Academic Press, London. Holm-Hansen, 0., S. Z. El-Sayed, G. A. Franceschini and R. Cuhel 1977. Primary production and the factors controlling phytoplankton growth in the southern ocean, In : Adaptations within Antarctic Ecosystems, pp. 11-50. G. A. Llano, Editor. Smithsonian Institution, Washington, D. C. C-32 Hopkins, T. 1971, Zooplankton standing crop in the Pacific sector of the Antarctic. Antarctic Res. Ser. , 17: 31+7-362. Hurley, A. 1976, Feeding behavior, food consumption, growth, and respiration of the squid Loligo opalescens raised in the laboratory. (NOAA) Fishery Bulletin 74: 176-182. Ivanov, B. 1970 Knox, G. A. 1968. On the biology of Antarctic krill, Eupbausia superba. Marine Biology, 7: 340-351. Tides and intertidal zones, Tn: Symposium on Antarctic Oceanography, Santiago, Chile, Septem- ber 13-16, 1966, pp 131-146. Scott Polar Research Institute, Cambridge. Laws , 1970 R. M, 1977. Antarctic marine ecosystems, In: Antarctic Ecology, Vol. 1, pp. 69-96. M. W. Holdgate , Editor. Academic Press, New York. Seals and whales of the southern ocean. Trans. R. Soc, London, 279: 81-96. Phil 1977b, Lockyer, C. 1976. Mackintosh, 1970. Mackintosh, 1974. The significance of vertebrates in the Antarc- tic marine ecosystem. In: Adaptations within Antarctic Ecosystems, pp. 411-438. G. A. Llano, ed. Smithsonian Institution, Washing- ton, D. C. M, Estimates of growth and energy budget for the sperm whale Physeter catodon . ACMRR/MM/SC/3 8 N. A. V/hales and krill in the Antarctic Ecology, Vol. twentieth century. In: 1, pp. 195-212. M. W. Holdgate, Editor. Academic Press, New York. N. A. and S, G, Brown Whales and whaling. Antarctic Map Folio Series, Folio 18: 2-4, C-33 Makarov, R. R. and V. V. Shevtsov 1972. Some problems in the distribution and biology of Antarctic krill . Translation: Israel pro- gram for scientific translation TT72-50077. Marr, J. W. S. 1962. The natural history and geography of the Antarc- tic krill iEuphausia superba Dana). Discovery Rep. , 23: 33-464. Moiseev, P. A. 1970. Some aspects of the commercial use of the krill resources of the Antarctic seas, I_n: Antarctic Ecology, Vol. 1, pp. 213-216. M. W. Holdgate , Editor. Academic Press, New York. Nemoto, T. 1959. Food of baleen whales with reference to whale movements. Sci. Rep. Whales Res. Inst., 14: 149-290. Oritsland, T. 1977. Food consumption of seals in the Antarctic pack ice. In: Adaptations within Antarctic Ecosys- tems, pp. 749-768, G. A. Llano, ed. Smithsonian Institution, Washington, D. C. Permitin, Yu . Ye., and M. I. Tarverdiyeva 1972. The food of some antarctic fish in the South Georgia area. J. Ichthyology, 12: 104-114. Prevost, J. In press. Population, biomass, and energy requirements of Antarctic birds - attempted synthesis. In: BIOMASS, Vol. 2, Background Papers. S. Z. El-Sayed, ed . Scientific Committee on Antarctic Research . Ryther, J. H. 1963. Geographic variations in productivity, In: The Sea, pp. 347-380. M. N. Hill, Editor. HiJiley Interscience , New York. Ven Heukelem, W. F. 1973. Growth and life span of Octopus cyanea (mollusca: Cephalopoda). J. Zool. Soc. London, 169: 299- 315. C-34 PERSONAL COMMUNICATIONS Hugh DeWitt Darling Center for Research, Teaching and Science University of Maine Walpole, Maine 04573 John A. Gulland Aquatic Resources and Survey and Evaluation Service Department of Fisheries Food and Agriculture Organization of the UN Via delle Terme di Caracalla 00100 Rome, Italy Robert J. Hofman Marine Mammal Commission 1625 "I" Street, NW Washington, D. C. 20006 Mary Alice McWhinnie Department of Biological Sciences DePaul University 1036 Belden Avenue Chicago, Illinois 60614 David J. Tranter Marine Ecosystems Group CSIRO Division of Fisheries and Oceanography PC Box 21 Cronulla, New South Wales, Australia APPENDIX F The Living Resources of the Southern Ocean: Krill Appendix F-i Southorn Ocean FieherieB Survey Prograunme GLO/SO/77/l THE LIVIKa RESOURCES OF THE SaTTHERN OCEAN lar Inigo Everson Consultant to the UNDP/FAO Southern Ocean Fislieries Survey Programme POOD AND AGRICULTURE ORGANIZATION OF TOE UNITED NATIONS UNITED NATIONS DEVEL0H1ENT PROGRAMME Rome, September 1977 F-ii The designations employed and the presentation of material in this publication do not imply the expression of any opinion whatsoever on the part of the Food and Agriculture Organization of the United Nations concerning m z o H G) > m TO q O -n n 5 n 5 o 7: ^ 0 m m o O o o o m z c I H o o > m 70 CI m z o m E.CRYSTALLOROPHIAS E. SUPER BA E. FRIGIDA E.TRIACANTHA E. VALLENTINI E. L0N5IR05TRIS E.LUCENS E.SIMILIS ANTARCTIC ZONE SUB-ANTARCTIC ZONE Fig. 6.1 Distribution of species of Euphausia in the surface waters of the Antarctic and sub-Antarctic zones (John 1937). The blacked-in portion of each column shows the normal range of that species, the entire column the possible range. Fig, 6.2 The et^hauBild Euphausia Buperba (3 om), the krill of Antarctic waters (Clarke and Herrinir 197 1) F-3 6.2 Distribution In broad terms the overall geographical distribution of ^ superba may be taken as being circunrpolar south of the Antarctic convergence (Marr 1962, Baker 1954) and is one of the dominant macroplankton organisms. (The dominant components of the macroplankton in the Pacific sector may be Salps and Coelenterates(Barkhatov et al. 197 3) although Mackintosh (1973) thinks most of the krill there are in the East Wind Drift and therefore in the Pack loe Zone). There are few records of Ej_ superba occurring north of the convergence. Mackintosh (1973) described krill north of South Georgia in a tongue of cold water, Sasaki et al. (1968) described a concentration at 45°S 145°E and Ball and Dunstan (1957) found fresh krill in the stomsich of a humpback whale caught off Queensland. Marr (1957) considers such oocurrances of minor importance. Within these very broad limits the density of krill is extremely variable. Marr (1962) describes the major concentrations as occurring in the East Wind Drift, Scotia Sea, Weddell Drift and South Georgia areas although Mackintosh (1973) has shown the existence of large concentrations elsewhere. As. a result of recent sampling Nemoto (1968) has shown the presence of large concentrations in the vicinity of the Kerguelen Gaussberg Ridge and at Longitude 150°W (see also map in Mackintosh 1973). This area of con- centration is also suggested hy Beklemishev (I96O, I96I) on the basis of whale catches in the region. Because krill concentrations tend to occur in certain clearly defined areas, it has been suggested that these may represent several self maintaining populations (implied by Mackintosh 1973) or even distinct races (Hakarov 1974). Clearly whether or not such a system is present will have direct effect on the way in which this resource is managed. In this review, the distribution 6f all life stages has been considered in order to lead to an overall understanding of the population structure. Distribution of eggs and early larvae In an analysis of the vast collection of material collected by Discovery Investigations, , Marr (I962) showed that E;_ superba spawns in the surface layers (althou^ Fraser (1936) con- sidered that spawning occurred on the bottom in the shelf zone). The eggs then sink and begin the processes of cleavage leading to hatching. The scarcity of cleaved eggs in the surface samples and the fact that Bargnann (1937) records spawning in aquaria but without subsequent cleavage implies that depth (probably pressure) may be necessary for development. Hatching is thou^t to occur in deep water and the resultant nauplii moult throu^ the metanaupliar stage as they rise through the water column to reach the surface as first calyptopes. This description of the cycle of events is clear from Marr' s (1962) analysis. What is not clear and which has been the subject of a great deal of study are the details of locality, time and conditions under which these events take place. Althou^ Marr (I962) recorded both gravid and spent females in the oceanic surface water, he considered that hatching in the oceanic deep water far away from the shelf was negligible. He did however consider that the early larvae were carried in the cold bottom water away from the continental shelf to undergo the developmental ascent in the oceanic deep water. This is shown diagrammatically in Fig. 6.3. In his analysis Marr considers very carefully all the evidence both for and against this explanation because it clearly has a great deal of bearing on the dynamics of distribution. In this respect the important factors are the time taken over the early development and the rates of flow of the cold bottom water and warm deep water. There is at present no direct information on early development rates for Ej_ superba. Marr assumed that Ej_ superba would take about half as long again as the 15 days Mefcanyctiphanes norvegica takes to reach the first calyptopis stage (p. 207). Mackintosh (1972) also considers three weeks to be a reasonable estimate although his analysis is based on much the same in- formation as Marr's. F-4 Fig. 6. 3 Vertical distribution and migration of the early development stages of krill on the shelf (left), and in deep water (right), (redrawn from Marr (1962)), Hatching in shallow water can give rise to the occurrence of nauplii and metamorphis un- usually close to the surface F-5 Little evidence is available on the rate of flow of the various deep water masses although Marr (p. 212) does ijive evidence that in places the Antarctic Bottom Water could flow north with considerable force. In a recent re-examination of the distribution of efcgs and larvae ,Voronina (1974) states that spawning occurs in oceanic waters (cites DolzhonkoT 1973 and Hakarov 1973) and also at South Georgia, a mar.fp-nal area of its distribution range (cites Mackintosh 1973 and Makarov 1972 )i/ and suggests that spawning probably occurs over most of the range of the species. Voronina suggests that the Antarctic Bottom Water rather than carrying the eggs and larvae does in fact form the physical lower limit for the sinking eggs. Successful development is assumed to be a direct fxinction of depth and a maximum depth of 1 8OO ra is ouggcsted as being the greatest vertical distance through which the larvae can migrate using the limited food resources in their yolk sacs. The 1 800 m boundary layer does encompass most of the localities in which early larvae have been found although Voronina does mention that some of the stations from which they were taken in the Scotia Sea fall outside this limit. Another criticism is that in many cases the larvae will not arrive in the surface water until late in the season after the peak of primary production. Adolescent and adult E^ superba have been shovm to be capable of feeding on detritus (Pavlov 1974) and there would seem no reason why the larvae should not do the same. In that case there is only an advantage in being at the surface during the primary production season. Thus althou^ the 1 8OO m boiindary layer may for the most part describe the northern limit of successful spawning this is probably a coincidence. In his analysis of the life cycle of E^ superba. Mackintosh (1972) to a great extent avoids the direct issue of oceanic versus shelf area spawning and concentrates on the dis- tribution of the subsequent developmental stages. However, the implication of his conclusions is that he, like Voronina, considers that spaiiming occurs over much of the ocean as well as the shelf area. He makes a strong case for the early larvae being distributed in areas where low sea temperatures extend to great depth (see also Marr, Fig, 81 ). The major areas he identifies as holding early larvae are the Ilhst Wind Drift, Bransfield Strait and the diver- gence zones he describes in the Scotia Sea and IVeddell Drift. Vertical distribution of later stages In a long and detailed analysis of the "Discovery" samples, Marr (1962) suggests that the major concentrations of _E^ superba occur in the top 100 m and within this stratum most fre- quently in the top 10 m. In his analysis Marr is careful to minimise error due to avoidance, a major problem with the 1 m .diameter and smaller nets from which his samples were obtained. Althou^ massed in the surface waters for most of the day and night Marr does sug.gest that there is a pattern of diurnal migration but was unable to specify the conditions under which it occurred. The supporting evidence that Marr cites, based on the depths at which whales and birds feed, is not particularly helpful in this context as it only confirms that krill occur near the surfsuie at some time during the day or night (a fact known from the net hauls). What is perhaps more important is that there is a clear indication (Marr 1962, Table 35) that adolescent and post larval krill are not present in si^ificant numbers in the V/arm Deep Water. Recent Russian research using echosounders has shown a tendency for sexually mature krill to undergo diurnal vertical migration in the top 80 m of the water column (Shevtsov and Makarov 1969), and Pavlov (1969| 1974) has obtained results indicating that this vertical migration 1/ Ifekarov (1972) refers to spawning in the frontal zone between Weddell and Bellingshausen water which would be oceanic. Mackintosh (1972) does not describe spavming as occurring near South Georgia but considers it possible although Marr (I962) considers that no successful spawning occurs in the region. F-6 is directly associated vrith feeding (although no indication is given of how widely applicable these observBtions are). VJhon the krill are feeding actively they rise to the surface and are dispersed. They come together as swarms when they are replete, stop feeding and then descend. Pavlov suggests that the onset of swann formation and the duration of the swarm is directly related to food availability. Nakaraura (1974) described a similar type of vertical migration although he does not link this to feeding pattern directly. Assuming that availa- bility of food is a major factor in controlling sv;arm formation it would be expected that swarms would occur more frequently during the summer when primary production is at its maximum than during the winter. Feeding studies by Pavlov (1971 » 1974) have shown that detritus plays an important part in the diet of E. superba which would allow than to feed all year round in deeper water. Although Pavlov~~(l969) describes the guts of krill cau^t at 300 m as being empty and there- fore not feeding, the fact that their main lipid store is not in the form of waxes (Bottino 1974) suggests they do not go v;ithouL food for long periods (see Sargeant and Lee 1975)* Also Mauchline and Fischer (19^9) suggest by comparison with other euphausiids that Ej_ auporba althou^ feeding on phytoplankton by preference will also feed on detritus and animal material. In addition to this information re.-^rding migration in the surface water Shevtsov and Makarov (1969) also describe significant concentrations at over 200 m depth although it is not clear whether these wore detected by echosounders or solely by fishing at predetermined depths. Confirmation of this observation may be inferred from Permitin's (1970) observation that krill forms an important componont in the diet of several demersal fish species in the shelf area (see also section on Fish in this paper). More recently significant deep water concentrations of krill have been identified in shelf areas of the Scotia Arc where there was no marked temperature discontiniiity (Fischer 1976), The fact that those recent observations conflict with those of Marr can be attributed to either a changing behaviour pattern by the krill or else due to the different techniques involved in estimation. Ifarr, in considering reasons for the disparity in the daytime and ni^t time catches finds some evidence that the difference is not due to a diurnal vertical migration beyond the normal range of the nets, but that it is more likely to be due to avoid- ance or the fact that subsurface swarms represent a poor target for obliquely hauled nets. The difference may also be due to the methods employed. There is clearly a need for stand- ardised sampling in conjunction with acoustic detection devices in order to quantify the krill concentrations in both the surface water and also in the intermediate water. Horizontal distribution of later stages Prom the time tiiat they arrive in the surface waters until they spawn two years or more later, a significant proportion of the krill must move, or be carried, to an area from which spawning will result in maintenamce of the stock. Over most of the Southern Ocean the surface water has a mainly easterly and to a lesser extent northerly set. The result of this is that krill which occur in that area vri.ll be naturally carried away northwards. Within the general area of the Ekst Wind and Weddell Drifts, which contain the major concentration, the distribution is very patchy. In addition to this the habit of E^ superba in forming swarms has made detailed distributional study liable to considerable error. Recent fishing expeditions have, however, identified concentrations regularly in clearly defined areas. These are in the area of mixing of the circumpolar and Weddell currents (Cershanovich and Lyubimova 1971) particularly to the north of the South Orkneys (Burukovskiiand Yaragov 1967), on the loe side (in terms of dominant wind and current) of islands and submarine ridges and also in areas of descending currents bordered by ascending currents (Elizarov 1971) and off South Georgia (e.g., Elizarov 1971, Makarov, Naumov and Shevtsov 1970, Bogdanov and Solyanik I970). In the Ea,ct V/ind Drift zone concentrations have been found around lOO^E (Hasu 1974) and from I30OE to 170°W ^Ozawa et al. I968). All of these observations were made during the summer months and also in areas of svjarms. The only major concentrations that occur outside the East VJind and Weddell Drift zones are reported from the north of the Ross F-7 ISO* W ISCTE ISO- Fig. 6.4 Principal concentra-tions of the Antarctic krill (Marr I962). F-8 Sea (Hodgson 1902f Marr 1^62) and the Ker.^uelen Gaussberg Ridge area (iJonoto I968) a region poorly sampled by Discovery Investigations (l6 stations total from November to Apri]). The catches by commercial type operations indicate a distribution very similar to that described by Marr (I962) on the basis of plankton net samples (his Fig. 5 and 6), (see Fig- ure 6. 4). The limited number of lar{;:o catches in xhe Nest Wind Drift zone can be attributed to one of two factors. Either krill in this sono do not tend to swann and althou^i present in large quantities do not occur in dense concentrations or their total quantity in the region is lov;er. Mackintosh (1973) sug/rests that krill concentrations in Indian Ocean sector of the Wcr-t Wind Drift are progressively /xa7,ed dovm by the vjhalcs as they migrate south. It is not however clear whether the whale migration south is maintained because of food require- ments. Theories of distribution control There are in the literature a wide variety of theories vihich attempt to explain the reasons for the occurrence of the various life stages around the Antarctic continent. None of these theories is totally proven. The more plausible will therefore be considered along with the major evidence that is available in support amd against. Perhaps the simplest explanation is that throughout their life history krill remain more or less in the same location (Marr 19'52). Mackintosh (1972) quotes an average surface flow rate in the V/est Wind and Weddell Lrifts of lO era/sec. which is within the range of swimming speeds determined by Semenov (1969) ^or adolescent and adult krill. Although! it is not known for how long krill could maintain this speed Marr (1962, p. 155) quotes from some notes made by E. R. Gunther who observed a swarm of krill swimming against an estimated /3 knot current for several hours. It is also possible for krill to be maintained in more or less the same locality by migrating between the warm deep water and surface water (they are known to occur in significant numbers down to 4OO m (Shevtsov and Makarov I969)) althouf^ Fischer (1976) has suggested that such a temperature discontinuity may represent a physical barrier. The eddying effect around islands and submarine rises could also, as suggested by Makarov (1972) and Khvatskiy (1972) hold krill in an area for a prolonged period. The remaining theories require that the krill are carried in a wide circulation pattern and the most frequently considered area from this point of view has been the Weddell Sea. Ruud (1932) basing his ideas on the circulation pattern of Meyer (1923) suggests that some larvae and adults get carried back to the hif^i latitudes in the surface stream. Marr (I962) whilst a.greeing that there is an undoubted circulation pattern of this typo believes that the circulation involves an area far further east and that in addition in order to complete the cycle the larvae would need to be carried south in the intermediate water during the developmental ascent. The easterly spreading of early larvae in the surface waters is clearly sliown in Marr's distribution maps (Fig. 64 onwards) which gives considerable weight to the greater part of this circulation theory. Makarov (1972 ) using recent information from the Scotia Sea has taken the theory still further and suggests that the main circulation pattern described above applies and the main stock is maintained by spaivning in the vicinity of the South Orkney Islands. He suggests that the larvae are carried around the Weddell Sea in a clock^irise direction to arrive back at the South Orkneys a year later. Some of these krill are carried around for a second year to spavm whilst the remainder drift out of this circulation pattern. Tliis theory requires confirmation of the existence of the recurvature current (see Kumagori and Yanagawa 1958 and also Fraser 1936, p I65) and also a better understanding of total circulation in the VJoddell Drift (see Deacon 1976). In an extension of this cyclonic circulation theory Makarov (1972) has suggested that a similar system to that in the V/eddell Zone could operate in the Ross Sea and Kcrguelen Gauss- berg ridge areas. Treshnikov (l97l) describes six cyclonic g>'res produced by prevailing cyclonic winds in coastal regions around t'ne continent. Thesn may represent areas of krill concentration as s u gested by Beklemishev (I96O). F-9 In his review of distributional control one of the mechanisms that Marr (1962) postulates is the possibility that a population of krill in the Bellingshausen Sea continually releases larvae and adolescents that seed the East Wind Drift. During the course of successive gener- ations this process is continued around the continent (see section on zooplankton in this paper). Such a theory althanf^ possible with water circulation pattern may not seem very likely in view of the known higher density of krill at the end of the cycle (Weddell Sea). However, this could be explained by the Weddell Zone itself .being a self maintaining aysteni as described above. I-Iarr also mentions the possibility that larvae could be carried from the Weddell to the Bellingliausen Sea via the Branafield Strait. Althougii there is no evi- dence for a water movement the full distance, there is evidence that water does enter the Bransfield Strait from the Weddell Sea (Clowes 1934 )• It has also been suggested (Everson 1976) that the Bransfield Strait itself might contain a self maintaining system. The only area containing large concentrations of krill that has not so far been consid- ered in this review is South Georgia. This has been considered as an area in which there is no successful spawning (Marr 1962) and as a result it is suggested that the krill population there has its origins in other regions. Because VJeddell Sea water has been considered to flow along the northeast coast of the island (Deacon 193?) and also because the Southern Scotia Sea has been shown to contain large concentrations of krill the Heddcll Sea is often considered as being the origin of most of the South Georgia Krill (Marr I962). Recent research has suggested that the frontal zone between IVeddell Drift and Wf^st Wind Drift now passes well to the south of South Georgia (Bogdanov _et al. 1969» see also section on Hydro- graphy in this paper). Under normal conditions therefore it would seem unlikely that Weddell Drift krill could be carried to South Georgia. Bogdanov and Solyanik (1970) in an attempt to explain how VJeddell Drift krill could get to South Georgia show a good correlation bet- ween whale catches and the mean annual air temperature at South Georgia and Laurie Islands (South Orkneys). They suggest that Weddell Drift surface water is carried to South Georgia as a result of wind action on the surface. Unfortunately the cold years (which should have produced largest krill concentrations and thus vjhales)were linked to low whale catches. Maslennikov (1972) considers that krill concentrations in the vicinity of South Georgia are present in the boundary region between a north westerly flowing coastal cvirrent and a south easterly flowing current further offshore. In years when this circulation pattern is ill defined, commercial krill catches have been poor (Maslennikov jet al. 1971 ). The impression is given in their paper that the water in this area is from the Antarctic circumpolar cui^ rent and that Weddell surface water is mixed in the manner described by Bogdanov and Solyanik (1970). It has also been suggested that the South Georgia Krill have their origins in the Bransfield Strait or Bellingshausen Sea area (discussed by Everson 1976) although there is only circumstantial evidence to support this idea. It is an interesting point regarding all of the proposed theories of ' distributional control that they all rely on fairly precise limitation in timing of development and -flow of water masses. This is in complete contrast to the mechanisms of distributional control proposed for other members of the macroplankton which in general rely on the southerly move- ment of the warm deep water and northerly movement of the surface water. (Mackintosh 1937» Foxton 1956, Voronina I968). For a mechanism such as this to be applied to krill it is clearly necessary to prove the existence of a seasonal vertical migration. Recent evidence has shown the existence of significant concentrations of krill dovm to 400m althou{^ as yet there is no year round coverage of observations to suggest that a seasonal pattern of verti- cal migration exists. Mention has already been made of the possibility that krill could feed in deeper water on detritus. The fact that the main lipid store is not waxes (Bottino 1974), indicating continuous feeding (Sargeant I975), would require the krill to be in deeper water during the winter. (With almost zero primary production in the surface water in winter the only food at that time is likely to be detritus at greater depth). Clearly there is a need for more research to identify which of these mechanisms are of major importance in maintaining the enormous standing stocks of krill. 1^-10 6. 3 Growth and Life Span Several authors have published curves of size at age for E^ superba (e.g., Ruud 1932, Bargnann 1945i Marr 1962) (Pig 6.5) which indicate a seasonal pattern of growth during a two year life span to a maximum size of about 6 cm. More recently Mackintosh (1972) has produced curves of "local apparent growth" based on the catches from specific regions. His analysis does not include a consideration of the origin of the krill S.t a given location but relies purely on the size of individuals at the time and place of capture. The simple two year life span has been questioned by several authors (Marr 19^2, Ivanov 1970, Hakarov 1971, Mackintosh 1972) who were all v;orking from infoimation from size frequency distributions. Recently Hakarov (1975) has demonstrated the ability of female Ej_ superba to spawn more than once thus indicating a longer life span than two years. I>larr (1962) because of the presence of a few specimens of abnormal size in his samples suggested that under certain conditions full sexual maturity and spawning may be delayed until the third year. This phenomenon has been reported in the related Ej_ triacantha (Baker 1959)* The degree to which the odd size group vra.s present in Marr' a samples was very small indeed, Hovjever, Ivanov (1970) with a much smaller number of samples detected an intermediate size group in a large proportion of samples from the Scotia Sea. He discussed this in detail sLnd concluded that the intermediate size group represented an additional year class. At the moment there is insufficient evidence available to conclusively prove that this intermediate size group does not represent an additional year class. Relevant information for both points of view will therefore be considered. At the same time that Ivanov was preparing his results Mackintosh (1972) was in the process of reanalysing in detail the Discovery material. As a result of his analyses he showed the presence of an intermediate size class in most of the areas of dense distribution. For those areas where there are sufficient samples Mackintosh has drawn an expected growth curve which shows the intermediate group as being midway between the previously accepted first and second year groups. Mackintosh argues that since the previously accepted one and two year classes follow on in a smooth curve with fast growth in spring and slow growth in winter, the cause of the intermediate size group must be something other than an additional year in the normal life liistory. On the assumption that the intermediate size fproup is real and not an artefact of sampling. Mackintosh considers the possibility of am early or late spawning producing the effect.' On balance he favours the theory that the intermediate group results from a second spawning later in the year (perhaps in May) but it should be emphasized that this conclusion is based on extrapolation from the start of the intermediate growth curve back for over a year. Makarov (1975? 1976) has shown tha^ a significant proportion of the population spawn more than once, first time spawners release their eggs during the later part of the summer and second time spawners in the early part of the summer. The release of eggs at more than one time during the season of maximtun primary production (and therefore growth) is likely to produce considerable blurring of the peaks in any size frequency distri- bution. Compared to _Ej_ triacamtha the size frequency distributions for E. superba show relatively indistinct peaks (see Baker 1959» Marr 1962, Barpjnann 1945t Ivanov 1970, etc.). Mackintosh (1972) detected a correlation between the presence of the intermediate group and an above average temperature in the preceding half year (but not earlier). Al thou^ it is unlikely that a warmer year would cause a second spawning, it is possible that a warm year could result in a greater disparity in the growth of larvae resulting from spawnings at dif- ferent times in the production season. Hakarov (1971 ) in a paper that also considers the conclusions of Ivanov (l970) is of the opinion that the extra size group does not represent an extra year class, but is a result of the mixing of krill from different sources. This suggestion is supported by the fact that the Bransfield Strait and South Geor,gia, areas of possible mixing, show the phenomenon quite frequently. Against the idea is the fact that the Bist Wind Drift zone also contains an F-11 60- 50- 40- 30- 20- 10- 0|N|D|J|F|M|A|M|J|J|A|S|01N|D|J|f|M|A|m]j|J|A|5|0|N|D|F|F|M -60 RUUD. 1932 FRASER, 1936 BARGMANN,I945 NEMOTO, 1939 -Q- POPULATION GROWTH (South Georgia) -•- 50 -40 -30 -20 -10 0|N|D|J|F|M|A|M|J|J|A|S|0|N|D|J|F|M|A|M|J|J1A|S|0|N1D|J|F|M Fig. 6, 5 Published growth cuarves of krill (redravm from Mackintosh (1972)) F-12 intemiediate group, but on infrequent occasions and since the IDast Wind Drift has the slowest growing krill of all areas, the mixing would have to be with krill growing more slowly even than these which is clearly impossible. Evidence on growth rate by other methods is at present inconclusive. Mackintosh (I967) Mcl^innie (1976) and Clarke (1976) maintained several individual adolescent and adult krill in experimental aquaria and studied growth over several moulte. In all cases growth incre- ment was very small althou^ both authors consider that their experiments were not a true reflection of growth in the natural state. The slow growth rate indicated by these experi- ments may indicate a slow growth rate in the wild. However, more sophistication in experi- mentation will be needed before this approach can be considered as giving a definitive answer to the problem. In an analysis of growth curves for a variety of euphausiids Mauohline and Fisher (I969) have calculated average daily growth rates. These are summarised in Table 6.1. Table 6.1 Calculated daily growth increment for a variety of euphausiid Crustacea (Mauchline and Fisher I969) Species Life Time (Days) Growth Increment (mg/day ) Thysanopoda acutifrons 730 0.356 Euphausia triacantha 730 0.246 '^eganyctiphajies norvegica (Iceland) 730 0.274 (Clyde) 730 0.427 (Cadiz) 365 0.383 rhysanoessa raschii 730 0.155 r. inermis (N. Iceland) 1 095 0.132 (3. Iceland) 730 0.101 r. longicaudata (Gulf Stream) 365 0.032 (Greenland) 365 0.028 P. longipes 0 730 0.100 $ 730 0,141 E. superba (Bargnann) O 730 0.849 $ 730 0.570 (Ruud) 730 0.808 (Nemoto) 730 1.340 (Mauchline) 730 0.822 The fact that the figures for E. superba are muc>i higher than those for any of the other species may indicate that the lifespan used as the base (two years) is too short. A greater lifespan would reduce the daily growtyi increment to nearer the values for the other species. F-13 A more detailed examination of the growth characteristics indicates that this hi^ figure for daily growth increment may not be totally unexpected. In his analysis of growth in E. triacantha, Baker (1959) suggests that the larger size of E^ superba is reached by a longer period of growth rather than by a faster average rate. Althou^ at first sight this conclu- sion appears to be at variance with the calculated rates of Mauchline and Pisher, whan consid- ered in relation to season the similarities become apparent. The percentage increase in length during each month for the two species Ej_ superba and JE^ triacantha has been plotted in Fig. 6.6 and the seasonal changes divided up into a series of growth stanzas leading to a first spawning. The changes occurring are tabulated in Table 6.2. The difference in the spawning season of the two species means that by September or October, Ej. triacantha larvae are twice the length of _Ej, superba larvae and it is only in the second growing season that Ej_ superba become larger than Ej_ triacantha. Bearing in mind that the _Ej. superba were only present for the last part of the productive season stanza C is effectively the first production season that the young Ej_ superba will experience. This is in effect the equivalent to stanza A for K. triacantha. Plotting growth during stanzas A and B for Ej. triacantha with those of E^ superba for stanzas G and D (Fig. 6.7) show how similar growth is for the two species during this period. Table 6.2. Subjective analysis of growth stanzas in Fig. 6. 7 leading to a first spawning in E. superba and Ej_ triacantha Growth Season Growth in Stanza E. triacantha E. superba A Summer Larvae present at Larvae present Hi^ primary production start of season rapid growth throu^- out season at end of season B Winter Very little primary production Slow growth rate Slow growth rate C Summer Moderate growth rate Past growth rate Hi^ primary production possibly due to de- throu^out most velopment of gonads of season later D Winter Slow growth rate Slow growth rate Veiy little primary leading to spawning production at end of period E SiiitrTier _ Slow growth. Hi^ primary production Build up to spawning condition Althou^ pre iding some evidence in support of the originally published growth curves this evidence shot ? mi^t be expected follow. Secondly, (normal for Ej_ triac not be considered conclusive for several reasons. Firstly althou^ it ^t similar species should grow in a similar way this does not necessarily account has been taken of growth by larvae resulting from the early tha) spawning in _E^ superba. The information related to growth in Ej_ superba is summarised in Table 6. 3. F-14 < I ^- < z m < (C o iij < Q. D V) hi UJ o z • o V) ■ < \ \ \ - 2 - 2 <. s \ \ • « a ^ o +^ M j3 CO •H o O a •H a P JS ^^ P. s +> S a o o O p. u c tt-^ E +• •H ca (d H V ■d ts -Cl t> +> 01 i-l 0) rH O V «] ■• +» ^^ -d X) ^t t> O 3 -p (d •• C! o r-H jq ;h - o - 2 - 5 I O o lO o -r o CD (o^ ) juauiajoui m3uai Am;uoi/^ o +» -EM •H V U 01 -P €) Jl ^ oj bo +^ pi v o ON & o s -p O O WlON C '- •H h O h V Cm « m a! tfl PI i o o ja Dl fl ^ ° -tf nJ O V ^ E o • ,^ -E) «M +> v^ C n) o Dl Ol o Id ■H -H P.WI bD •H P-15 < I 1- < z CD < a u Ml 4 a. -) q: w ^- UJ UJ - o - < - z - < - s - z - o XI +» XI 0 a ^ - < T o -r o in -r o T o (% ) ;u3maj3ui q}3u9i Aimuop\[ F-16 Table 6.3. Summary of infonnation related to life epsm in EuphauBJa euperba Observationa Evidence for 2 years Evidence for more than to first spawning 2 years to first spawning Size frecfuency rjialysis "Smooth" curve "Intermediate" size group of Bargnann, etc. "Intermediate" Does not fit amticipated Additional peak or peaks size group smooth curve in many size frequency distributions in siunmer Growth rate based on Summer growth pattern Overall growth rate appears two years to first similar to E. triacantha high spawning Laboratory growth Probably not valid for Measured growth rate very experiments "natural" conditions slow Recent work (reviewed in the section on Reproduction )has shown that a significant pro- portion of the population spawn for a second time althou^ the growth increment in this additional year is only small. The information that is available althou^ far from conclusive indicates that the growth pattern originally described by Ruud (1932) and Bargnann (1945) is probably correct. There is however considerable doubt about this and the possibility that the intermediate size group does represent an additional year class has neither been conclusively proved nor disproved. 6.4 Reproduction A good general description of the reproductive biology of euphausiids is given by Mauchline and Fisher (1969), whilst detailed specific information covering the main aspects of reproductive biology is available in Bargnann (1937) and Praser (1936). The topics of particular interest in this review are spawning season, fecundity and the phenomenon of repeated spawning. Spawning season It is now generally accepted that spawning by Ej. superba occurs at the surface (see Marr 1972, Pig. 30, Voronina 1974, Makarov 1974) during the summer months. In a brief review of published information on the precise spawning period Mackintosh (1972) suggests that it is at its peak in February and March althou^ gravid and freshly spent females were found from November to April. In general he describes the 3i)awning season as being earlier in the northern end of the range than nearer the continent and gives the following "best estimates" of mean hatching and spawning dates (Table 6.4). The dates (Table 6.4) were calculated to provide the origins of growth curves from the different regions and are estimates based on the known spavming dates in the Weddell aind i^st V/ind Drift zones. Thus, although they are presented within fairly broad limits and are subject to local annual variation, they are useful at this stage as a first approximation. F-17 Table 6.4' (From Mackintosh 1972) Mean dates for major spawning and hatching of Ej_ superba Locality Spawning date Hatching date 3outh Georgia-' 25 January 3 February Northern Weddell Drift 31 January 9 February Scotia Sea 20 February 1 March Central and Southern V/eddell Drift 5 Iferch 14 March Bransfield Strait 11 March 20 March East Wind Drift 20 March 29 March V/orking on samples collected in the vicinity of the South Orkney Islands (equivalent to Mackintosh's Scotia Sea area) Makarov (1973) describes spawning as occurring at the end of February and that the timing of the peak is very much dependent on temperature. In warm years he suggests the sea ice clears earlier thus allowing an earlier start to the growing season with a resultant earlier spawning. The main purpose of Mackintosh's analysis of spawning date was aimed at providing an origin for the gi'owth curves for results from different areas. He was therefore only con- cerned with the mean date of spawning and took little account of the several month variation in the observed dates on which eggs were found. Marr (1962) in considering the overall spawning season in the northern or Weddell zone did not consider spawning as a short-term outburst, as had been suggested by Ruud (1932), but suggested it occurred over the period November to March. The recent work of Makarov (1975 » 1976) has indicated that individuals spawning for the first time do so late in the season (February, March) whilst those which live for a further year spawn for a second time during the November, December, January period. The release of eggs during most months of the production season will almost certainly result in a broad size frequency distribution for each year class (if year classes can be identi- fied). This in turn could also result in the production of an intermediate size group (see section on growth) as has already been discussed. Clearly it is only when definitive spawn- ing periods in given locations are identified and the development of the subsequent size groups are followed that it will be possible to subdivide the broad size frequency distri- butions and thus help in unravelling the growth story. The infonnation reviewed by Hauchline and Fisher (1969) indicates that euphausiids gen- erally spawn in the spring or early summer and that by spawning at the end of the summer E. superba is an exception. It is however possible that originally the spring breeding season now identified by Makarov may have been the norm and that the late summer spawning has only recently evolved. No matter what the sequence of evolution it is probable that the two spawning periods represent a fail safe device in maintaining a hif^ biomass. Fecundity Estimates of fecundity are few and include a wide range of values as a result of diffei^ ent techniques in estimation. These are reviewed by Mauchline and Fisher (I969) who consider 1/ (N.B. There is very little evidence of successful spawning in the vicinity of South Georgia). F-18 that for euphausiids in general the best estimation technique is that based on the calculated nvunber of egga whose volume is equal to half the mature ovary and that the ripe ovary occupies about 10^ of the body volume. Makarov (1975) states that the eggs released amount to 34-39?^ of the body wei^t in E. superba which, assuming that wei^t is proportional to volume and that eggs have a volume of 1 x lO"'^ ml, indicates a very much higher fecundity. This estimate is in closer agreement with the results of Nauraov (1962) who counted ripening eggs present in the ovary. The estimates are compared in Table 6.5. The fi(^e of over 11 000 detarmined by Bargnann (1937) is probably an estimate of all the eggs at all stages present in the ovary rather than an estimate of the number that would actually be released. This figure has there- fore not been included in the tabulated results. Table 6.5. Fecundity of Euphausia superba. Data from Mauchline and Fisher (l969)t Makarov (1975)1 Haumov ( 1962) and Hemoto et al (in press). Body Length Body Volume No. of eggs ( 1 X 10~4m (each)) Equal to 505J Equal to Counted ovary 34-39?^ Body Volume mm ml mg Mauchline and Fisher Makarov Nemoto Naximov 36 2362 38 2655 40 3007 41 0.62 310 2110-2420 3590 1 43 43.5 851 3502(2222-7792) 3520, 3320 44 2960] 45 3480] 1.0 3400-3900 1.6 800 5440-6240 46.7 927 7146(2482-14086) 8370(3889-13091) 50.3 1507 It is not clear what is the reason for the enonnous variation in the estimated values of fecvmdity. The egg size noted by Naumov (I962) in the ovaries of the individuals he examined would indicate that the eggs released represent nearly 60^ of the body volume. The release of such a large proportion of the body volume as eggs just prior to the winter season seems unlikely if the same individual is going to spawn a second time in the following spring. On the other hand the general figures derived by Mauchline and Fisher (1969)1 as they point out, are not always correct and in this context it may be that Ej_ superba is an exception. There is therefore a clear need to identify what proportion of the eggs present in the ovary are released and also to generally improve the refinement of these estimations. 6.5. Swarming The habit of Ej_ superba of forming dense aggregations has been well known for a long time and because of the concentration of individuals within the swarms they are often thou^t of as ideal for exploitation. V/hat is not clear however is the proportion of the total krill population that is present in the swarms nor the absolute density. The former point is vital to resource management and the latter to rational fishing. F-19 In this context a swairo can be defined as a dense aggregation of individuals moving ha]>- moniously and is probably the type of aggregation that will give a strong indication on an echosounder. A swarm may therefore be considered as the extreme case in terms of co-ordinated movement of individuals in a group. The other extreme would be independently random movements by individuals. Between these two extremes there almost certainly exist concentrations that are intennediate in terras of aggregation and organization. Swarming has already been discussed briefly in the section on zooplankton where it was suggested that swarming was a result of some social behaviour pattern (as yet unidentified) working in areas of hif^ average concentration (such as discussed in the section on krill distribution). The section which follows is a review of information on swarm formation and disintegration. Marr (1962) concluded on the basis of analysis of plankton net samples that from hatching onwards the number of krill outside of swarms. was negligible. Although Marr was well aware of the maximum density krill attained when swarming his definition of a swarm was not nearly as restricted as that given above. Other authors have been more evasive on the point only mentioning swarms without making any firm commept on the proportion of the total population present in swarms. The observa- tions of Pavlov (1969) and Shust (I969) indicate that swarming is a transient phenomenon related to food availability. These observations were based on the state of the gut in freshly cau^t animals, a method also used by Nemoto (1968). According to Pavlov when phyto- plankton is abundant the krill spend part of the day actively feeding in the surface water until they are replete at which time they aggregate into swarms and sink. In one region in which Pavlov made his observations this cycle occurred twice in 24 hours. The cycle of swarm- ing and dispersion was repeated in other areas although without vertical migration; Pavlov attributes this difference to one of size; adolescent krill, he suggests, migrate vertically whereas adult krill do not. Shevtsov and Makarov (l969)» however, note that in the course of vertical migration the adult krill tended to be deeper and in fact on occasions formed a separate layer (detected by echosounder) beneath the adolescent krill; they make no comment on the causes of swarming. Nakaraura (1973) identified several type:5 of krill aggregation and related their presence and formation to light intensity. He found the densest svrarms at the surface on the darkest ni^ts and when the ligti't intensity increased the density decreased as the animals migrated down. The commonest pattern of aggregation that he identified was a layer generally within the range 30 to 100 m depth or about 20 m above the thermocline. The results of Ozawa _et al. (1968) conflict with these observations and indicate surface aggregations in daytime; they were hovrever a result solely of visual observations. Shust (I969) considered that the level of illumination (cloudiness) of the environmental factors he measured (wind, sea state, atmos- pheric pressure) was the most important although he considered feeding also to be of major importance. The suggestions of Ruud (1932), Beklemishev (I96O) and Bogdanov (1974) that turbulence and eddy currents are responsible for the formation of svrarms is probably not valid on the above definition of a swarm but more likely applies to the formation of high average abundance. Marr (1962) and Naumov (l962a) suggested that some swarms were in part spawning concentrations. Althouf^ there does not appear to be any conclusive evidence (no catches have knowingly been made of krill in the act of spawning) Makarov (1973) has shovm the presence of concentrations of ripe individuals during the spawning period; he does not, hoifevcr, describe the concentra- tions as swarms as defined above. It has also been suggested that swarming is a result of increased buoyancy due to the increase in lipid content during the summer (SurukovBkii 1967 )• Presumably this would be expected to have the effect of concentrating the krill into LanfTtiuir cells as described by Stavn (1971) for Daphnia although there is no evidence to suggest that for krill this does in fact occur. F-20 The major factors that have been positively identified as being associated with swarming are therefore light and feeding activity. Reactions of planktonic animals to light have been well documented by Gushing (1951)1 Stasenko (I967) and Burukovskii (196?) describe krill being attracted to a red light source but moving away from a strong white light source while Ivanov (1969) has drawn attention to the luminescence of krill in swarms. Fiirther information is clearly needed to identify a possible wide variety of reactions by krill to light. The suggestion by Pavlov (I969) that krill swarm as part of a feeding cycle implies dependence to a great extent on food availability - the more food that is available the more rapidly the animals become replete and swarm. As mentioned earlier a direct result would be that swarms would tend to appear during the early summer and be most frequent in abundance during the summer peak of primary production. Hardy and Gunther (1935) showed that in the South Georgia area the densest concentrations of phytoplankton were not coincident with the largest concentrations of krill (or other zooplankton) (see Table 4«l)» This general pattern is confirmed by Avilov et al. (I969) who found the largest concen- trations of krill in areas of the Scotia Sea where the mean biomass of phytoplankton was I-5 ml/m^. In areas where the mean biomass of phytoplankton was less than O.5 ml/m-^ or more than 5 ml/m they found few krill concentrations. This observation is not confirmed by Shust (1969) who reports numerous krill swarms in areas where the phytoplankton density varied from 0.1 to 8 ml/fflJ, He does however relate duration and size of swarm to phytoplankton standing crop — swarms persisting longer in areas of high phytoplankton standing crop. It is possible that the inverse relationship reported by Hardy and Gunther (1935) and Avilov et al. (I969) is due to grazing effect whereby there is a time lag before the herbivores eat down the phy- toplankton (Gushing et al. I963 and Gushing 1975)* Tha direct relationship in that case wovild be expected during the initial phytoplankton bloom in areas where overwintering herbivorous zooplankton concentrations occur. Although this explanation is not proven in this instancei it does fit the facts and therefore indicates that studies investigating this interaction, and thus by implication other factors that might control krill raicrodistribution, should take accovmt of short-term changes. Swarm Size Estimates of swarm size have been made by both visual observation and analysis of echo- sounder records. Marr (I962) published a series of drawings of the horizontal outline of several swarms that were observed near to the surface. These swarms were all quite amorphous in shape and varied in size from a few to several hundred metres across and were continually changing shape in an amoeboid fashion. Nemoto et_ al. (in press) describe surface swarms as being generally round or oval in shape and that some changed shape as a result of wind stress becoming more oval with the greatest axis at right angles to the wind direction. A comparison of swarm size between areas is made by Nemoto et al. (in press) who con- sidered that swarms in the Scotia Sea area were on average smaller than those in the Queen Maud Land area. In the vertical plane swann size has been mapped using echosounders. Certain of the small swarms are so dense that it is impossible to d^efine the vertical size. However, where density within the swarm is lower, authors have mapped the vertical distribution and size of swarms. In an analysis of this type, Shevtsov and Makarov (I969) describe various patterns in the changing shape of swarms observed in the Scotia Sea. From their observations it appears that there are a few common patterns throiighout the area. The following types of vertical movement and configuration are described in Table 6.6. Formation of two layers (Table 6.6) during the day and amalgamation at night with a tendency for adult krill to be at the deeper levels seem fairly standard. A much greater vertical movement was observed "by Mohr (1976) who, using an echosounder, followed a concen- tration of krill in the vicinity of the South Sandwich Islands for 6 days in April I976. At midnight the krill were massed in the top 20 m but by O6OO hours they had migrated to a depth of 70-110 m, to slowly migrate back to the surface during the following 10 to 12 hours. Since all these observations were made in a fairly brief period of time during one season, more information will be required to amplify these observations and conclusions. F-21 Table 6.6 Information on the vertical range of krill swarms (From inforroation in Shevteov and Itokarov) Locality Depth Ranee (m) of: Composition and other comment Upper layer Lower layer Day Nifcht Day Nirfif Northern Weddell Sea 2 Layers day time; one layer ni£^t, nearer surface. More adults with increasing depth Northeast of S. Orkneys (a) (b) 10-30 15-30 (10-40) 30-60 40-70 One layer at night. Catch mainly adults. One layer at night. Mainly juveniles near surface. Mainly adults at depth (200-400 m). S. Orkney S. Sandwich 10-15 10-15 Mostly juveniles West of Sandwich 10-20 20-45 40-80 Surface swarms also. Few mature krill. N.E. of South Georgia 15-40 50-100 Few adults S.W. of South Georgia 40-60 40-60 Pew adults, no vertical migration. Swarm density Several authors have published estimates of the density in terms of ntunber and weight of krill within swanns. These are summarised in Table 6.7. Most of the estimates (Table 6.7) of 3v;arm density vjere made by means of acoustic fish- finders in conjunction with actual catches. There are clearly several sources of possible error in this type of estimation. Although the ochosounder may give some indication of the vertical extent of the swarm it ivill give no inaication of its lateral size; also, it is often difficult to confirm that a particular acoustic indication resulted in a particular catch. The two estimates given by Marr (I962) are based on separate obsei^ations. The figure of one per cubic 'inch is based on the density estimated by an observer looking vertically down into a swarm from above. The figure of one per ei^t cubic inches was calculated from a single lateral haul in a large swarm (the net was towed for 36 sec at 1-J kt through the swarm). Assuming a mean wei^it of one gramme per individual, this represents a density of about 6kg/m^. F-22 Table 6.? Estimates of krill density in sv/arras Density Reference Numerical By Wei gilt I per in. 3 1 per 8 in.^ Marr 1962 10 - 16 kg/m3 up to 15 kg/m3 or more Moiseev 1970 Makarov et al. 1970 generally up to 5 kg/m3 max. 6-33 kg/m3 IJemoto and Nasu 1975 ?000 - 8000/m3* fax. 40 OOO/ra^ mean 1.5 kg/m3 Nemoto et al, (in press) « Absolute range 521-49,153 Ag a first approximation therefore the acoustic estimates largely confirm those from plankton hauls. From the point of undertaking fishing operations it is clearly possible to detect krill concentrations using echosounders. Tlie magnitude of any diurnal vertical move- ment is likely to be of little consequence in catching technique. However, greater sophisti- cation is required to quantify echo sounder indications in order to use them for estimation of standing stock or as "ground truth" for estimation by satellite imagery. 6.6 Food and Feeding An extensive study of the food organisms found in the stomachs of a large sample of Euphausia guperba was made by Darkley (1940). The species list that he gives is largely confirmed by the observations of Ilart (both lists are discussed at length by Marr 1962, pp. 172-176). iniile there are local differences in the occurrence of the various species (see also Ilustedt 1958, Nemoto I968, Kawamura, in press) the diatoms clearly predominate from these analyses. The preponderance of diatom remains is generally thou^t to result from the relative indigestibility of the siliceous frustules and therefore probably does not represent the true situation (see discussions by Marr I962 and Mauchline and Fisher 1969) and althouf^ the feeding appendages are adapted to a herbivorous diet it seems clear that, when necessary, representatives of other groups can be taken. Mauchline and Fisher (I969) list the following groups as being represented in the diet:- Algae, DiatcMS, Dinoflagellates, Tintinnids, Radiolarians, Foraminifera and Crustacea. In addition, Pavlov (1971) increases the list to include detritus and in extreme cases cannibalism. As already mentioned vdth regard to distribution detritus may well represent a major part of the diet in winter although there are no results to confirm this. F-23 The only information available on diurnal feeding cycles is that of Pavlov (1969) who related the occurrence of svjarms, feeding and food availability, (see section 6.5). It is not however clear to what extent thcoe results are typical of the Southern Ocean as a whole or how they vary seasonally. Since the presence of certain diatoms in the guts of krill can have adverse effect on the quality of products made from them there is clearly a need for more study in this field. 6.7 Production and Biomass The extreme variability in the density of krill caused mainly by its swarming habit has presented enormous problems in estimating biomass and production. There are quite a large number of papers giving estimates calculated both by direct and indirect means and these will be considered along with the major sources of error. Based on visual observations from the deck of niiS William Scoresby and assuming a den- sity of one individual per cubic inch in the top yard of the water column, Marr (I962) estimated a mean biomass of 2.5 g /m'^ for krill. In the same paper he recalculates the biomass using the results of Hoyerdahl (1932) for mean weight and gets a figure of 29-28 g/m^. The possible error in these estimates is enormous since the area under consideration is 500 yards on either side of the ship's track and swarms are assumed to be one yard deep. These rough calculations presented with great reserve, as I>Iarr puts it, may be» he thinksirepre— sentative of the Sist Wind and Weddell Drift zones. Assioming that this area is about half that described by Mackintosh and Brown (1956) as south of the Antarctic convergence the total biomass is 521 million tons (based on 29-28 g/m^) or 44-5 million tons (based on 2.5 g/m^). Itorr, basing his reasoning on the probable consumption of krill by whales is of the opinion that the hi^er figure is the more reasonable. This is the only direct estimate I have so far come across that is supported by calculation and yet even in this case the figures are presented within very broad limits since they are calculated by extrapolation from a rela- tively very small area that was at the time seen to be unusual. In an analysis based on echosounder survey and fishing catches, Neraoto _et al. (in press) have estimated standing stock as being from 0.5 kg/m-^ to 30 kg/m-* althou^ they do not indi- cate how these figures apply to the total Southern Ocean. Makarov and Shevtsov (1972) quote a range of 953 to 1 350 million tons for the bicmass of krill (abstract only seen) while Gulland (197O) estimated the total zooplajikton biomass to be about 10 g dry wei^t per square metre or I50 million tons for the whole Antarctic and assumes that 50^^ of this is krill. Based on Gulland' s estimates the krill biomass is there- fore 75 million tons dry weight, or perhaps 750 million tons wet wei^t. The figure of ^0% for the proportion of krill in the zooplankton is open to question as Voronina and Naumov (1968) found that Euphausiids made up only 7-6^ of their zoopleinkton catches. However, since their nets may well have missed a significant proportion of the Euphausiid population this figure cannot be taken as firm even though it is based on a large number of observations. To estimate production from these biomass figures, a conversion rate of 1:1 has been suggested (Gulland 1970, Hempel 1970) based on a mean life span of one year. Allen (1971 ) has questioned this ratio and by comparison with mortality rates in other Euphausiids (there being no infoimation available for Ej_ superba) he suggests that the production is between 1.8 and 2.1 times the biomass with the lower figure the more likely. On this basis the krill production is 940 million ton/year from Marr's figures and 1 350 million ton/year based on Gulland' s figure. Indirect estimates of biomass and production have been made based on both higher and lower trophic levels. First of all considering estimates based on primary production, Hempel (1970) quotes results from localities all round the Antarctic continent for annual carbon fixation of 43 g C/m2 (Currie I964), 84 g c/m2 (El-Sayed 1967) and 100 g c/m^ (Ryther I963) and suggests that since the measurements tended to be in the richer areas a figure of 50 g c/m2 is a reasonable approximation. Taking a conversion ratio of carbon to fresh wei^t F-24 of 1t10 primaxy production is estimated as 10 000 does not appear in the literature any firm figure by Euphausiids although a 10^/ figure has been use (1974) estimate that in growing to I.5 g an indiv gives a conversion ratio of lt40, but since this achieve a maximum size the mean population figure ally used \J . On the basis that krill represents ous zooplankton, the production figures ralculate million ton/year (Oulland 1970), There for the conversion rate of phytoplankton d (Oulland I970) . Chekunova and Rynkova idual consumes 60 g of phytoplankton. This is an average figure for an individual to may well be nearer the 1:10 ratio gener- between 10 and 50^ of the total herbivor- d are given below; Conversion Ratio Phytoplankton: Krill 1:10 1:40 Krill as Proportion of Herbivorous zooplankton 505^ 500 125 la/, 100 25 Estimates of krill production (millions of tons per annum) calculated on the basis of total primary production of 10 000 million ton per annum. Calculations of the amount of krill eaten by whales have been made by several authors and more recently the same calculation has been made based on the estimated initial whale stocks to give an estimate of "surplus" krill. Independent estimates of krill consumed by whales are given in Table 6.8. Laws (lnp?8BB)has also estimated the amount of krill consumed by Crabeater, Leopard, Ross and Fur Seals at 53 920 thousand tons each year of which about 99^^. is taken by the Crabeater Seal . Estimates of krill conuumption by birds have been made by Provost (in press) and Croxall (pers. comm.). Both agree that the major consumer is the Adelie Penguin (PygOBcelis adelie)which it is estimated takes about two-thirds of the 14-7 to ?0.3 million tons total eaten by birds. There are no estimates in the literature for krill consumption by other groups such as fish and Brpiid although it is quite likely that either or both could be major consumers. Laws (in pre^)eBtimates that sperm whales consume annually 4.6 million tons of scjuid which, assuming they feed predominantly on krill and assuming a ten to one conversion ratio, would account for 46 million tons of krill. The same calculation applied to the estimates based on orfuid eaten by the total initial whale stocks would account for 102 million tons of krill per annum. 1/ In their paper Chekunova and Rynkova give a detailed breakdown of food consumed at monthly intervals during the lifespan. In the absence of realistic size frequency information it is not possible to produce a better conversion ratio. F-25 Table 6.8 Estimates of Annual Consumption of Krill by Whales Author Krill eaten by ini- tial whale stock Ton X 10^ Krill eaten by present whale stock Ton X 106 Potential "Surplus" Ton X 10^ Marr (1962) iS — — Studentskiy (19^7) - 270 - Kasahara (I967) 24 - 36 (Pin Whale) Zenkovich (l970) 150 - - Mackintosh (1970) 120 - 170 (10)»»»* 100 - 150 Hempel (1970) 45- 60 - - Gulland (l970) >50 - - Nemoto (1970)» 77 - - Doi (1973)** 200 - - l^yubimova et al. (1973) (800 - 5000 )#»* - - Ohfflura 250 (40)*»»» 100 - 200 taws (1977) - - 153 Laws &n press) 1^ n Ml Results where all the factors in the calculation have been given are \inderlined » Cited by Nemoto and Nasu 1975 but reference not given ♦* Cited by Nanoto and Nasu 1975 *** May refer to total krill biomass *•♦* Calculated from his results N.B. (There is a much longer list of references quoting krill constiraption by whales but those seen are derived from those tabulated). The total estimated prcluction based on consumption by predators is summarized in Table 6.9. Clearly, in spite of the enciroous uncertainty attached to these estimates (Table 6.9) the total amount of krill consumed by predators is large,, thus confirming the estimates by other methods. P-2'^ Table 6.9. Estimated consumption of krlll by maior predators \J (for detailed breakdovm set- relevant consumer section) Estimated consumption of krill prior to whaling million ton/year Estimated consumption now million ton/year Whales Seals Birds Squid Pish 190 (?) (?) (?) (?) 43 64 15 - 20 (100 ?) (?) TOTAL >190 > 200 6.8 Eyploitation Annual catch Exploratory fishing for krill has been in progress now for several years. Solyanik (i960) obtained a small catch of krill using a small pelagic net from a v;haling ship. Catches from the subsequent krill fishing expeditions are shoim in tables 6.10, 6.11 and 6.12. In addition to these exploratory fishing expeditions there are catches reported in FAO Yearbook of Fishery Statistics Vol. 40 (PAO 1976) that probably refer to krill ("Marine Crustacea, unspecified " taken from areas adjacent to the Fisheries Statistics Antarctic regions). These figures, together with actual krill catches recorded in the Yearbook, are set out in Table 6.13. 1/ A single estimate of standing stock of between I.5 and 5 milliard tons is given by Bogdanov (1974)« Since this figure was calculated on the basis of consumption by whales and other consumers it probably refers to prodviction rather than biomass. Even allowing for the product ion/biomass factor proposed by Allen (l97l) this figure is still enormous compared to all other estimates of the resource. In the same paper Bogdanov estimates the standing stock in the Southern Scotia Sea in I967 as 5 million tons, half of which is in the zone of mixing between V/eddell Sea and circumpolar water. Also, he estimates that at South Georgia the .stock is 0.7 million tons. These estimates were made directly and although the precise method f nployed was not stated it was probably using echosounders and commercial catch data. F-27 Table 6.10 Exploratory fishing by vessels of the USSR Vessels Season Catch (tons) Reference Muksun 61/62 (70)^ Burukovskii and Yaragov (I967) Muksun 63/64 Stasenko (I967) Aoademik Knipovich) 64/65 185 Nemoto and Nasu (1975) Orehova, Obdorsk ) 121 Aoademik Knipovich 66/67 •p Aoademik Knipovich 67/68 >140 Ivanov (1970) Table 6.11. Exploratory fishing by Japanese vessels Vessels Season Catch (tone .) Reference Chiyoda-Maru 72/73 59 Nemoto and Nasu (1975) Taishin-Maru No. 11 73/74 646 Nemoto and Nasu (1975) Taishin-I'Iaru and Abo— Iteru 74/75 1 140 1 460 Anon (1976) Exploratory Vessel) 75/76 Commercial Vessel ) 5 000 Anon (1976) Planned Catoh 76/77 (10 000) Anon (1976) Table 6.12. Exploratory fishing by other nationalities Nationality of Expedition Season Catch (tons) Reference Chile 74/75 75/76 60 Anon 1975 West Qermany 75/76 400 Anon 1976 a (2 vessels) (77/78) Poland (2 vessels) 75/76 ? Anon 1976 b Other Asian Countries 76/77 ? Anon 1977 Norway 76/77 Small Anon 1977 a 1 / The total catch in his tables is only 23 tons F-28 Table 6.13. Catches of krill in area 58, and of unspecified Marine Crustacea by certain countries in areas 41 1 47 1 51» 57 snd 87 metric tons Country^-~-^__^^ 1969/70 1970/71 1971/72 1972/73 1973/74 1974/75 Krill Japan U. Marine Crust USSR Chile 100 1900 1300 1200 2100 4400 7400 2600 643 4412 821 1081 6965 821 In addition to these figures McWhinnie (1974) and El-Sayed (l975)i estimate that a total of 200 000 tons of krill have been taken per season in recent years. This figure is not supported by further evidence and it probably refers to total landings of krill and fish (gee section on fish exploitation). From the foregoing it is evident that the present fishery, taking as it does about 20 000 tone per year, is still only having minimal effect on the resource. However, all of the nations presently involved in exploratory fishing are capable of mounting quite a con- siderable effort and thus increase the size of the fishery very quickly. Catch Rates Development of a fishery based on a novel resource means that exploratory fishing oper- ations may well meet with only limitp 01 s o (D ■!-• «H s H^ O Dl ^ 8rH S -d f>s a w <0 « 4) a .H ^ .^ >-< 4* ^-^ +■ +^ ■H 0 d 01 rH n* -d to o a o cd ig V is rH m o ■r1 bS 01 d w 4) d >J +> C fn -P -2 to •H d o c O H o c g rH •H O -H ^ ^ a^ 0 d +J Td M >> . o d W bo r-f s, 0) .H 0) .s +» bD •rl o c; r-l P «H •rl d -P •H +^ rl n3 § • o to •H o +^ •r^ ^ i +^ +» d x! d o a w O Q) S° S 0) s 4J 4) ■d 4) ■d d 4J -d &. t>j bo O -P O 0 o 2 t. •H U •H ■r4 ■rl •d U s . ^ rt nJ (3 a rH •H += •H •rl 9 ,; ■6^ •rl r. ^ s o 1 1 ^^ 6>^ a ;j 4) "d • 0) ^< ^^ •H O •H H •rl 4) -rl r-\ tfl to n) a, a li-. W -H W S W ««-' m E id O o +> M ^ ■(-> S d •rl ■r^ (4 a n> d D bo o •H ^ 0 +5 o ■A 8 ^ +» m ■d •rl rH .% ?0 •H (3 H* rH rH ^ 0) j:: rH 8 •rl rH to bO O iH O f^ tlH > U nJ •H (1> 4) o ^< >; (0 o a «H ^ O +J t-H rM a 65 rH •H S rH rH •H rH rH @1 M rH rH O (0 0) 1 1 d C) ^^ d <§ 1 S fe 1 •H rH 10 0) •^ <^ 4) 4) t>J tJ hD S>5 4) e -d 2 ^ rH HJ :a o § rH "S ra +> tJ nJ n ro •rH o bo c ^ ■d 4) ^t be a O d §s 0) t^ li 3 4) bo nJ Q ■H p. to u d M !>jpq . rt . <" •H o •H d ^ rH . bD O bD P^ tH «M Pi -H nJ 4) hD rt -H ci o< 4) d a, CO a 2 a M rt O -Q (fl -d 4) -p ;^ O -rl rH -d +^ 4) bD M bD^ s 0) O 4) •H (l> •H O •d •S"' d 4) g 0) o HJ •H «"> J2 d JU rH nJ > U 4) •H •H O- u i 5 nJ 8 s •rU .• 8 ^ m P. m d CO o •H a p< ^. y. s •—X O. 4) ^— N *'— N o ^ m tn nJ nJ ,Q M^-^ (13 rO c a o ^^^^ ^,_^ ,^ o • * +^ n) -Q o +> r-l C\J 8 8 «H Z V. E. V. V,V' = Value per unit weight of prey and predators respeotiirely E,E' = Exploitation rate of prey and predators respectively k = Conversion ratio prey to predator w,w = Mean wei^t of individual items eaten and in the catches respectively The summation is over all prey species. In the form that the equation is written here the greatest value would result from harvesting the predator. With only sli^t rearrangement this general ecfuation may be applied to the krill/whales type situation. In this case there is only one prey species but several predators. Using the same notation except that the summation is over the different predator species (whales, seals, etc.) the equation then becomes: n I V! EI K . W, > V E v7 7 i 1 1 i Implicit in this equation is the fact that, as Gullajid pointed out, the efficiency of utilization (K) should be greater than the product of the relative value, the relative exploitation and the probable growth of the prey. In this equation a lot clearly depends on the relation between V and V and K. Gulland (1974) suggeststhat the overall conversion efficiency of krill to v/hales is verj-- low and of the order of 2/^ and suggests that if the problem facing the world were of more food at all costs, and the choice were between 1.7 million tons of whales, and 100 million tons of krill, there would be no doubt that preference would have to be given to harvesting krill. Clearly this represents an extreme case which in reality seems unlikely to occur. Betv;een these two extremes there are an infinite number of potential answers particularly if the other exploit- able resources are included into the analysis. The problem then reduces to one of making management decisions to utilize and at the same time safeguard the resources. An alternative indirect method of harvesting krill has been proposed by Joyner _et al. (1974) who suggest that the West Wind Drift is a suitable environment for salmon (see also Hardy 1975, Joyner 1976). On the basis of published information on seasonal sea temperature and the distribution of krill they consider that salm6n fry released from Southern South America could search out and feed on krill in the Antarctic. Because of the known fast growth rate of salmon they estimate that a salmon fishery of several million tons could be established in the Tierra del Fuego area. They point out the obvious advantage of such a system that since the salmon will probably return to their area of release the expense involved in fishing will be minimal. However, from the inf oiTnation tliey give it seems un- likely that the salmon will feed on krill since the main concentrations are south of the area regarded as seasonally acceptable to salmon (Fig. 6.8 which is derived from information in Joyner et al; 1974) • F-36 150 170 170 Areas of seasonally acceptable temperature to salmon t:^\'^v^\^ Summer distribution of krill Fig. 6. 8 The northern limit of krill distribution (see Fig. 6.1) compared with that of the range of surface temperatures acceptable to salmon (sheided area) (from Joyn«r e_t al . , 1974 ) APPENDIX G The Living Resources of the Southern Ocean Fish G-i Southern Ocean FisherieB Survey Progranime GLO/SO/77/l THE LIVING RESOURCES OF THE SOUTHERN OCEAN ^ Inigo Everson Consultant to the UNDP/FAO Southern Ocean Fisheries Survey Programme FOOD AND AGRICULTURE ORGANIZATION OF TOE UNITED NATIONS UNITED NATIONS DEVELOFT^IT PROGRAMME Rome, September 1977 G-ii The designations employed and the presentation of material in this publication do not imply the expression of any opinion whatsoever on the part of the Food and Agriculture Organization of the United Nations concerning ',ij -^ Continents and Islands RAJA GEORGIANA RAJA MURRAYII RAJA EATONII Fig. 8.1 (a) Distribution of main finh species in the Southern Ocean G-3 30°W 30° E «0°w 50° W 60°* 60° E 90° E 100° E II0°E 20° E I30°E ltO°E I50°E I60°E 170° E +++++++ Antarctic Convergence Ice shelves Continents and Islands MICROIVIESISTIUS AUSTRALIS Presumed yeor round distribution Known area for summer migration Fig. 8.1 (b) Distribution of main fish species in the Southern Ocean G-4 20°* 20° E 30°W 30° E 40° E 50°W 80° E 90° E 00° E I50°E I70°W iBo l?0°E +++++++ Antarctic Convergence Ice shelves Continents and Islonds ^^^^^ NOTOTHENIA MAGELLANICA Fig. 8. 1(c) Distribution of main fir.li species in tfift Southern Ccean G-5 20° w JO°W^ 40° W 50° w 70°W 80° W 90° w lOCW (I0°W I20°w^ rjo"*' >\. ^, V\ vr • 40° S J 120° E +-»-++V+++^ I50°E I40°E y I50°E +++++++ Antorclic Convergence Ife shelves Continents and Islonds NOTOTHENIA .■ RIFRONS Fig. 8.1 (d) DlBtribution of main fioh apecieB in the Southern Ocean G-6 80° E 100° E + + + + 4- 4- + Antriirlir Convprqence Ice shplvP'; Conlinpnl"; nnd l<;l(inils NOTOTHENIA NEGLECTA NOTOTHENIA CORIICEPS Fig. 8.1 (e) Dinlribution of main finh species in the Southern Ocean G-7 60° W 60° E 80°* go'E JO'E +++++++ ^^iii^ Antorctic ConvPrqence ice shelves Conlinenis and islonrls A NOTOTHENIA ROSSII B NOTOTHENIA ROSSII MARMORATA Fig. 8.1 (f) DietriViution of mnln finti npocies in the Southern Ocean G-8 10° w 60° W 70° W 80° W 80° E 100° e + ♦ + + 4- + + ^iSO^-u^ Antarctic Convergence Ice "Shelves Continents ond Islands . DISSOSTICHUS MflWSONI ( Large " specimens) the species are Known to occuc all oround ttie continent B DISSOSTICHUS ELEGINOIDES Flg.8»1 (g) Distribution of main fieh species in the Southern Ocean G-9 ioo°w V^ 100° E +++++++ flntorclic Convergence ICP 'Shelves Contmenlfi ond Islands CHAMPSOCEPHALUS GUNNARI Fig. 8.1 (h) Diotribution of main fioh species in the Southern Ocean G-10 20° W 20° E JO°W 50°W 70° E B0°W II0°W 90° E lOI7»t 120° E I40»E I50°E I60°E +++++++ Antorctic Convergence Ice shelves Continents and Islands CHANNICHTHYS RHINOCERATUS _..~ PSEUDOCHAENICHTHYS GEORGIANUS Fig. 8.1 (i) Distribution of main fish species in the Southern Ocean G-11 10° w 20° E 30°W WE 80°W 90° E +++++++ Antarctic Convergence Ice shelves Continents ond Islonds CHAENOCEPHALUS ACERATUS CHAENOCEPHALUS BOUVETENSIS Fig. 8.1 (j) Distribution of main fish species in the Southern Ocean G-12 40°W 60° w 20°W SO»W, 10° w 80° W 90° W o'^'' u^^.^. v\ 0 ° <)0°S 15° S ++ + + *AAii 50°S7~ ^BOUV 60°sy~~ ro°s/ ^ yK^'X-'' '■■f'l* 20° E 30° E + + + H 40°E A ANTARCTICA \i \. ^^SS sf/j! veo'E KERGC ^70»E "to 1 + %\ 1 + #1 \ Si'-Jl f 7 — T 90° E 100° E ' 1I0°E l?0°W ^ X I50°W / 50° S 1 45°S 40° S ^ I70°W 180' +++Vf+4-+^ 120° E I30°E I50°E 170° E +++++++ Antarctic Convergence Ice shelves Continents and Islands CHIONODRACO Sp Fig. 8.1 (k) Distribution of main fish species in the Southern Ocean C— 13 Table 8.1. Fish species of potential commercial importance in the Southern Ocean, Group Species Common Name Rajidae Ra.ia fceorriana Rt murrayi R. oatonii Gadidae MicromesiGtiuG auntralia Southern Blue Whiting or Southern PoutaGoou Morluclidae Merluccius hubbsii Patagonian Hake Nototheniidae Notothenia fabborif rons N. coriiceps N. nc^lccta N. roGGii roscii N. ronnii mannorata N. ma,n;ollanica DisBotichus mawsoni D. elef^noiden Ploura/xramma antarcticum Marbled Notothenia Antarctic Tooth Fish Patagonian Tooth Fish Channichthyidae Champsocophalus f^mnari Channichth,vG rli.inoceratus Psoudochacnichth.yn f:oorp:ianuG ChaonoccphaluG sp. Chionodraco sp. Although there are several species of hake in the Southern Heminphero, Morluccius hubbsii is the only npecies to have been roporbod from Antarctic waters (Hilcheyev I967). In this single instance fish v;oro assumed to have migrated into the Antarctic to feed on krill. This migration is not conaintent with the shorcvfard aummer migration pattern in Patagonia described by Hart (1946), a movement confirmed by the observations of Ciechomski and Weiss (1974) who found that M. hubbsii spawned in shallow water during the summer. The restriction of the distribution of many species to the continental shelf zone has probably resulted in the establishment of distinct stocks (even if not genetically at least as far as fishery management is concerned). Species where this is thouf^t to be the case along with the defined localities are listed in Table 8.3. G-14 Table 8.2. Distribution of Antarctic Fiah (See also Fig 8.1 a - k) SpocieB * Habitat and Mode of Life Depth Range Reference Ra.ia p;eor/Tiana Demersal, South Georgia Shelf also from a submerged elevation between S. Orkney and S. Sandwich Island (-1.44 to -1.47°C) Demersal, Kerguelen Demersal, Kerguelen 180-830 20-60 30 1, 2, 1 1 3 R. murra.yii R. eatonii Micromesistius australis In the Antarctic. Reports indicate this opeciee is generally pelagic in the vicinity of the continental shelf. Has been caught in bottom trawls althouf^ larger catches have been made with pelagic trawls. 200-650 10-70 2, 4, 5, 6 Merluccius hubbsii Reported on only one occasion Pelagic, assumed migrated to Scotia Sea from Patagonia 7 Notothcnia /^bberifronn Demersal in shelf area of Scotia Arc Demersal, Shelf area Kerguelen, Crozet Demersal in Shelf area of Scotia Arc and around continent Juveniles demersal in shallow water Adults demersal/pelagic in shelf area Originally considered coantal spec- ies living on kelp, now known to be pelagic krill feeder Mainly pelagic in open ocean Mainly pelagic Open ocean pelagic although often associated with Continental Shelf 5-350 0-200 0-200 0-30 0-400 0-80 20-220 70-800 1 1 1 8, 9 1 10 10 11, 1, 14 N. coriiceps N. nerilecta N. roGsii roosii and ) N. roGsii marraorata ) N. maKellajiica DioDootichus mawsoni D. elofcinoidog Pleurap:rarama antarcticum Champnoccphalua /?^innari Pelagic/demersal in Shelf area Demersal Demersal/pelagic in Shelf area Demersal DemorBal 0-450 0-140 0-270 5-350 0-800 12, 1 13, 1 12 12, 1 1 ChannichtKvo rhinocora- tUG PDoudochaonichth,yo Koor- fTianura ChacaioccphaluB ap. Chionodraco ap. G-15 References for Table 8.2. 1. DeWitt 2. Permitin 3. Bigelow and Schroeder 4. Merrett 5. Baoalaev and Petukhov 6. Shuntov 7. Mikheyev 8. 01 sen 9. Hureau 10. Yukhov 11. DeHitt and Hopkins 12. Olsen 13. Hureau 14. LiYubimova et al. 1971 1969 1965 1963 1969 1971 1967 1954 1970 1970, 1971 (a), 1972 (in Press) 1955 1966 1973 Table 8.3. Antarctic Fish Species for which there is positive evidence for more than one discrete managoraent stock. Species Stock Localities Reference Hicromesistius australis (a) Scotia Sea, probably oouthem limit of Patagonian population. (b) Campbell Plateau (a) and (b) Related subspecies on morphological grounds (a) and (b) Not considered of sub- specific status but considered isolated This species has also been reported from S. W. Indian Ocean Inada and Nakajnura 1975 Shpak 1975 FAO Motothenia rosail (a) Kcrcuelcn Crozet group con- sidered of subspocific status N. rossii rossii, Richardson (b) Scotia Arc. considered of sub- specific status N. rossii raarmorata, Fischer Nybelin 1947 Nybelin 1947 G-16 g,^ oize and Growth In the Antnrctic ichthyofauna small epeoieB ire dominant, over half the epeciee do not attain a length of 25 cm and few species attain a length of over half a metre (Andriaehev 1965). It is an interesting point that most of the larger species belong to the family Channichthyidae, a gro\ip possessing neither Auictional erythrocytes nor any respiratory pigment. Growth rates are, with few exceptions, slow as might be expected in an environment of continually low temperatures. The limited information that is available is set out below, Raja sp. The South Georgia skate, Raja georgian\TS is one of the largest Antarctic fishes. Permit in (I969) gives the siae/weight composition for 31 fish from a large sample caught in the vicinity of South Georgia (Table O.4). No information ie available on growth rate. Table 8.4. Length/Weight information for Itaja georgianus. Total Length (cm) Wt.(g) No. 20 - 30 146 3 30 - 40 - - 40 - 50 860 1 50 - 60 1 780 3 60 - 70 2 680 11 70 - 80 3 800 2 00 - 90 6 015 4 90 - 100 7 690 4 100 - 110 10 850 2 110 - 120 13 700 1 WicromesistiuB austral io Information on growth is only available for this species from Patagonia and the Scotia Sea areas (Table 8.5.). Table 8.5. Length (cm) against Age results for Micromcsiatius austral is Locality AGB GROUP Reference I II in IV V VI VII vm n X S.Orkney la. o" Q 40 45/47 45/46 40/49 48/49 Mikheyev I967 S.Orkney Is. (f 2 h 46 45 48 47 49 40 49 50 50 50 52 51 53 53 (>5 Ghubnikov, et al. 1969 Patagonia CT Q 44 44 45 46 46 47 47 48 49 49 50 52 55 50 Shubnikov, et al. 1969 Patagonia cT 13 21 29 37 Inada and Nakamura 1975 New Zealand cT 13 22 29 47 Inada and Nakamura 1975 G-17 The results of Inada and Nakamura (1975) refer to fork length and the remainder (although this is not stated) probably refer to total length. The difference in size at age between results of Inada and Nakamura and those of the Russian workers is almost certainly due to gear selectivity although the possibility of its being due to interpretation of the otolith rings cannot be ruled out. Merluccius hubbsii Information relating size to age for this species was published by Mikheyev (1967) and is Buminarised in Table 8.6, Table 8.6. Size at age information for Merluccius hubbsii in the Scotia Sea Length (cm) Weight (g) Age (yr) 34 - 35 260 (?) 21 - 27 - - 28 - 35 - 2 34 - 35 - 2 Notothenia neglecta Results from two localities, Terre Adelie (Hureau 1970) and Signy Island, South Orkneys (Everson I970) are set out in Table 8.7. Table 8.7. Size at age information for Notothina neglecta Age Group Length (cm) Weight (g) TA 0 Signy TA^ Signy TA 5 Signy TAq. Signy V 22.0 16.5(6) 22.5 311 107 318 VI 18.2(22) 24.0 18.5(7) 24.6 141 401 147 415 VII 21.4(22) 25.9 20.4(20) 26.4 225 502 190 513 VIII 24.6(10) 27.5 22.3(18) 28.0 338 607 240 612 IX 27.8(8) 29.0 24.2(9) 29.4 480 716 316 705 X 31.0(6) 30.5 26.2(5) 30.4 670 827 387 794 XI 34. 2» 31.8 28.1(7) 31.6 890 939 500 879 XII 37.4* 33.0 30.0(2) 32.5 1 157 1 048 620 960 XIII 40.6(1) 34.] 31.9* 33.4 1 485 1 150 760 1 034 XIV 35.0 34.1 1 245 1 101 XV 35:8 34.7 1 337 1 161 ( ) « Figures in parentheses are sample size. * " Calculated value (The data from Signy Island are calculated values based on examination of 2 000 otoliths) G-IR Hureau (1970) tried to fit a Bertallarify curve to his results from Terre Adelie but could not obtain a realistic estimate for L oo . He therefore derived the follovjing altei^ native equations. L^ = 3.2 (t) - 1.0 (5 t > 6) \ = 1.925 (t) + 6.9 (Cf t > 5) A Bertallanfy curve was fitted to the results from the Slgny Island fish and the constants are set out in Table 8.0. Table 8.8. Constants for the Bertallanfy curve derived for N. neglecta at Signy Island K Leo W«3 t 0 cf 0.091 0.129 ^15. 5 39.5 2 744 1 728 -1.7 -o.n Notothenia rossii Size at age has been studied at Kerguflen (llureau 1970) and also at South Georgia (Olsen 1954, Crisp and Carrick 1975). These results are Bummarised in Tables 8.9 and 8.10. Table 8.9. Notothenia roasii rosaii from Kf^rguelen (Hureau ^970). AOEGHOUP 1 I II III JV V VI VII VIII IX X Mean Standard Length Q 11.2 (12) 17.1 (17) 23.5 {62) 32.1 (23) 39.3 (9) 50.0 (1) /lO.O (3) 55.0 (2) 53.0 (2) 60.0 (2) 57.0* 64. o»^ 61.0 (1) 67X)» 63. 0» Mean Weight (g) 41 131 293 75? 1200 1963 2800 3815 ( ) = Kigures in parentheses are sample size. * = Estimateii value G-19 Table 8.10. Constants of the Bertallanfy curve for N. roasil rosali from Kerguelen. K L CO t 0 9 0.13 0.13 80 90 -1.69 -0.62 Table 8.11. Size at age information for Notothenia roosii marmorata Tram South Georgia (Olsen I954) Age Group I II Ill IV V VI VII VIII IX X XI XII XIII XIV Len^h (cm) 0 7 - 22 28 32 44 51 59 61 64 67 70 72 72 50 57 59 62 64 65 68 69 Bertallanfy curves were fitted to the data for South Georgia with the constanta that are eet out in Table 8.12 (Everson 1970). Table 8.12. Constants of the Bertallanfy curve applied to data of Olsen (1954) for Notothenia rossii from South Georgia. Age Group K L CO t 0 I - V V - XIV 0.15 0.29 75 75 0.1 3 Tlie change in the growth pattern had previously been deocribed by Olsen (1954) as being coincident to, and therefore probably a result of, the offshore migration of the fish. The age determination methods employed by Olaen and Hureau in their studies were based mainly on otolith examination although they both examined scales as a check. Crisp and Carrick (1975)) from examination of the scales from fish of known size cau^t at Leith Harbour, South Georgia, were able to corroborate Olsen's size at age estimates both by direct comparison and by back calculation based on the ratio of scald length to total length corrected for allometry. Their rosulto confirm the growth pattern described by Olsen for the first few years and also indicate that this techniqute is of great use in obtaining the maximum amount of information from small samples. Notothenia magellanica The only published information on growth of this species is from fish oau^t at Kerguelen (Hureau 1970). The relevant information is given in Table 6.13. G-20 Tatle 8.13. Size at ago information for Notothenia magellanica at Kerguelen. A g e G r 0 u P I II III IV V VI VII Length (cm) Weight (g) N 12.1 55 4 18.0 178 18 23.4 374 23 26.6 565 11 30.3 642 11 33.1 886 5 36.5 1 176 5 Hureau has fitted a Bertallanfy growth curve with the following constants to these results: K = 0.26; L CO c 40cm; t ■= 0.4yr. o DissootichuB mawconi and DiBSOBtichus eloginoidoa Yukhov (l971a) hao described a series of specimens obtained from sperm whale stomachs the size;; of which are set out in Table 8. 14. Table 8.14. Length/vioight information for DisoostichuG mawooni (Data from Yukhov 1971a) Length (cm) Weight 0 (kg) Weight ^ (kg) 121-130 24.6 131-140 29.5 29.8 141-150 38.9 37.2 151-160 53.0 49-0 161-170 55.0 171-180 70.0 In addition Calhaem and Chris toff el (I969) Cive the following information: Species Length (cm) Weight (kg) Notes DioooatichuB mawooni ' 147 125 30 (10) 5p. in Dominion Museum, Wellington, New Zealand Taken from seal There is no information available on age determination or size at age for either species althou^ Yukhov (l971b) has described the otoliths in detail for taxonomic purposes. G-21 Pleuivi.'^yainma ajitarcticum No information on gi-owth is available for this species. They are reported to grow to 30-35 cm total len^rth (300 g) (Lyubimova et al . , 1973) Champsoc ophalus r\mnari Olsen (1955) published information on the size at age for this species at South Georgia and this is summarised in Table 8. 15. Table 8.I5. Length at age information for Champooccphalua f^unnari data from Olsen (1955). Age Group III IV V VI VII VIII IX X XI XII Mean Length (cm) N 26.8 18 32.5 11 34.9 24 36.3 13 36.7 10 38.2 5 41 1 0 41 1 42 2 % 26.6 29 32.1 24 34.2 21 36.5 10 37.6 9 39.5 4 40.5 2 40 1 41 3 0 Channichth.YS rhinoceratus Huroau (1966) has published information on size at age for this species and this is summarised in Table 8.16. Table. 8. 16. Size at age information for ChannichthyG rhinoceratus Age G r 0 u P II III IV V IV VII VIII IX X XI XII Mean Length (cm) 14.8 22.3 26.0 34.0 37.0 39.7 41.0 42.5 — — 47.5 Mean Height (g) 35 150 220 435 510 550 635 750 — — 1 330 N 0 2 6 7 8 4 3 1 0 0 1 Poeudochannichthyg georglanua Detailed information of ago and aizo for tliis species is lacking although Olsen (1955) given the following information based on examination of the vertebrae of formalin fixed epeclmena. First year 6 - 10 cm Second year 17 - 23 cm 2-3 yearn old 27 - 33 A nlnfije individual ( + ) of 50 cm was aged 8-9 yoars. G-22 Chaenocephalus aceratus The results of Olsen (1955) for this species at South Georgia are svimraarised in Table 8.17. Table 8. 17. Information on length at age for Chaenocephalus aceratus Age Group IX X XI XII XIII XIV XV XVI XVII Mean Length (era) d 55 52 51 51 52 52 50 N 1 5 6 3 3 2 1 0 0 21 $ 62 63 57 59 65 6? 65 N 1 3 1 0 3 2 0 1 2 13 Several v;orkers have published equations relating length to total weight for Antai-ctic fish (Table 8.l8). Unfortunately a simple standard equation does not seem to be applicable although all are variations on the simple cubic relationship intuitively expected. Table 8.18. Length/v/eij^it relationships for Antarctic fish Species Relationship Locality Reference Haja .'eortaanus W = (0.21L)3 South Georgia Calculated from results of Permitin (1969) Micromecisticus see (0.21L - 1.3)^ Footnote (l) Scotia Sea Estimated from re- sults of Shubnikov (1969) australis Notothenia neglecta W = W = 0.029(L*)3 1 0.028(L*)3 ^ Signy Island Signy Island Everson 1970 log W = 2.92 log L*-4 45 Terre Adelie Hureau I97O N. roasii rossii log W =2.76 log L*-4 06 KerguQlen Hureau I97O N. roHsii ^marmorata W = 0.029IA-85 South Georgia Crisp & Carrick 1975 N. ma^^ellanica log W = 2.87 log L*-4 20 Kerguelen Hureau I97O Pseudochacnichthys W = (0.213L)3 Scotia Sea Dubrovskaya and Makarov I969 geor,"p.anun Chacnoc ophalus W = w - (0.189L)3 47.8 X 10-6l2-75 Scotia Sea Kerguelen Dubrovskaya and Makarov I969 Hureau I966 aceratus Channichthys rhinoceratuc (1) This equation does not fit the results of Dubrovskaya and Makarov (1969) who worked on fish from the same hauls. An approximate conversion ratio based on their results is W - (0.2L)3 W = Total Wei^t (g) L « Total Length (cm) L*= Standard Length (cm) G-23 S.^j Reproduction There arc in the literature few published accountc of seaoonal reproductive cycles but the detail that io presented in them has meant that the main patterns identified can be used to provide a reasonable description for those Bpocies for which only limited information is available. Hureau (1964) used the factor "Relative Gonad Size" (conad wei^t as a perccntag^e of total weight) from monthly samples to doncribe the gross cycle in the gonads of TromatomuB bemacchii. From those results and the results from several other species of Nototlieniid (Hureau 1970, Evorson 1970) it is possible to outline a poneralised pattern for the ovarian cycle. This in indicated in Fifj. 0.2, and with only sli/^jit adjustment of the (y) axes could be applied to most of the Notothoniids invostip;ated (Notothenia ncKlecta (from 3cotia Arc), N. ronsi 1, U. ma.^iollanica, Trcmatomus bomacchii, T. hanaonl ). The exceptions are Notothonia ne.'^'loc ta at Terra Adclit- which produces small diameter ova and therefore does not have such a clearly defined cycle and N_j_ cyanobrancha for which thfere are two spawning periods (those maturing for the first time spavm in January and those that have spawned previously subse- quently spawn in May, (llureau 1970)). Information on egg size, fecundity and repawning period for those species of potential commercial importance is given in Table 8.I9 and this has been combined with infonnation for all Antarctic species to describe major spaiming ty^ies (Table 8.20). It is a characteristic of Antarctic fish to produce large yolky eggs and this has been discussed by several authors (Harsliall 1953, Andriashev 1965« Permitin 1971, 1973). It is generally considered that large yolky eggs spavmod in the Autumn or early winter months will result in the production of post larvae in the Spring coincident ^^^. th the phy toplankton bloom. These large eg,gs are also generally considered to be pelagic (or benthopelagic in some cases). Tlio release of large eggs in the spring by Trematomus sp. may indicate a much slower develop- ment rate of the eggs and early larvae although there is no proof of this suggestion. Table 8.I9. Egg size, fecundity and spavming period for Antarctic fish Species !';gg dia. mm Fecimdity X 103 Spavming Period Reference MicromoDiotius Spring Hart 1946, Weiss 1974 autj trails Merluccius hubboii O.0-O.9(P) Deoomber Ciechomski and Weir33 1973 Notothenia 130 - 100 Jul -Aug Permitin &. Sil'yanova 1971 gibborifrons N. ncglocta 3.0(D) 10 - 30 (12/g Total Wei^t) May Ever son 1970 3 10.1 - 1/1.7 (8-16/g Total Weight) Permitin & Sil'yanova 1971 N. roncii rossii 1.2 3(D) ?0-30(l3-20/g Total Weight) 46-53(l5-17/g Total ^ Weight) January May Hureau I97O Hureau 1970 N, rossii marmorata N. magollanica 4, 8(D) 1.2(P) 20 - 120 60 -70 (50-58/g Total Weight) Apr-May Feb/March Pertnitin * Sil'yanova 1971 Hureau I97O G-24 16- 14- 12 0) m •g IO- CS B o 0) > 2- RELATIVE GONAD SIZE OVA DIAMETER r t -1 1 r 4 9 6 10 II T- 12 Months since last spawning T— — 1 I 2 3« 3.4 -Jl -3.0 2.8 -2.S -24 -2.2 a a 2.0 U (U 0) -i.« a •a > O hl4. 1.2 f-lO o.a o« 0 4 h08 Flg>8.2 Oenoralized pattern of changes in the ovary of Antarctic notothenide in the pre-epavming period G-2 5 A Boriea of subjeotivo stages in the gonad cycle of Nototheniids has been described by Elverson (in press) based on the ovarian and testis cycles in Uotothenia neglect a (Tables 8.21, 8.22). Table 8.21 Stages in ovarian maturation of Nototheniids based on the cycle in Notothenia neglect a Maturity Stage Description 1 . Immature Ovaries small, firm, no eggs visible to naked eye 2. Maturing Virgin Ovaries about 74 length of body cavity, firm, full of small eggs 3. Developing" Ovaries large, contain eggs of two sizes 4 . Gravid Ovary large. Vfhen opened, large ova spill out 5* Spent Ovary flaccid, contains a few large and many small ova Table 8.22 Stages in the testis cycle of Nototheniids based on the cycle in Notothenia neglecta Stage State Description 1. Immature Testis very small and translucent, lying close to vertebral column 2. Developing Testis small (about 1^ total weight), white and convoluted 3. Developed Testis large, white and convoluted. No milt produced when pressed or cut 4. Ripe Testis large, opalescent white. Drops of milt produced under pressure or when cut 5. Spent Testis much smaller and flabbier than stage 4, dirty white in colour G-26 Table 8.I9. (continued) Species Egfi diem, mm Fecundity X 103 Spavming Period Reference DisGostichus 4-4. 3(D) (Nov/Dec)l Yukhov 1971a mciMGoni Champs occphalus 1.9-2.2 3.4-4 2.6 4-23 S. Georgia 8 S. Orkneys April May 01 sen 1955 L<3nnherg I905 Permitin 1973 Permitin 1973 Charapgoc ephalua Champoocephalua Chaenocephalus 4.7 3.9 S. Orkneys S. Georgia (June) Mar/Apr Everaon I96O Permitin 1973 Channichth.ys 4.5 6-14 Feb/Mar Hureau I966 rhinocoratus Pscudochacnichth.yo 4 5.&-IO.9 Mar /Apr Permitin 1973 Chionodraco 3.7-5.0 2.5-4 Mar/Apr Formitin 1973 Notea: 1. CompariGon of the process of final ovarian maturation in this species with N. ncglecta from Signy Island suggests that spaiiming occurs in the middle of winter and not Nov/Dec. P " Pelagic Eggs D = Demersal Eggs Table 8.20. Egg size and spawning time for Antarctic fish Spawning Period Egg Size (mm) Species Spring (Nov. Dec.) Summer (Jan. Feb. ) Autumn (Mar. -June) 2.5 - 3.5 1 2-4 TromatomuG bemacchii, T. hansoni Notothonia nGglocta (Terra Adelie) N. magellanica Notothenia nudifrons (jl. anpustifrone) iT. ncnlcx^ta (iJcotia Arc) N. ror.Gii R, kempi N. gibberifrons N. larneni (Dinoootichuo mawsoni) Champs ocophalun gunnari ChaenooeplialuG acoratus Pscudochaenichth.YG georgianun Chionodraco sp. Channichthys rhinocoratua Hureau I966, 1970} EKrerson 1970} rormitin 1971,1973; Koysner et alj_, I974 G-27 The eeaeonal cycle in the ovary of all fish species eo far studied, whereby there is a steady build up of oggs all of the same size, indicates that H].)awning Beaeons of limited duration are normal. The result of this is that it is to be expected that just prior to and during spawning the mature fiah will be congregated together. That this occurs is undoubtedly the case as concentrations of mature fish have frequently been caught on th« continental shelf of some of the island groups. The approximate localities and environmental characteristics of these localities are summarised in Table. 8t23» It should be borne in mind that the list. only includes localities where presjawning or spawning fish have been caught and obviously does not include all spawning areas. It is probably fair to assume that spaw- ning concentrations will occur in similar circumstances to those described in the table over most of the range of each individual species. Table 8.23. Sjiawning localities and environmental characteristics of spav-ming groiinds. Species Spawning locality liivironmenta] Characteristics References Depth(m] Temp. C Microinesistius Patagon ia S.G. S.O. ,(SG)(SS)Shelf S.G., Shelf Kerguelen Shelf S.G. Shelf S.G. Shelf S.G. Shelf 115-600 115-750 10-450 120-350 250-350 100-125 115-350 115-350 0.7 to -0.1 to - -0.4 to - 0.7 to 1.8 to 1.2 to 1.4 to 1.2 to 1.7 -0.4 -1.8 1.7 2.0 1.6 1.7 1-5 Southern jiart of Patagonian - Falkland Shelf Shelf Shelf Shelf North and East side of Island Rocky shelf SE of Island In Fjords and shallow parts of Shelf Shelf to N,S &E Shelf to N,S &E Shubnikov et al. (1969) Permitin and Sil 'yanova (1971) Everson (1970 Permitin and Sil 'yanova (1971) Keysner et al. (1974) Permitin (1973) Permitin (1973) Permitin (1973) austral is Notothenia gibberifrons N. neglecta N. rossii ma^-morata M . ro Bs i i rossii Champsocephalus gunnari Pseudochaen irh- thys georgianuB Chaenocephalus (S.G.: South Georgia; S.O,: .".oul.h Orkneys; S.S.: South Shetlands) Mention has already been made of the slow growtli rate of Antarctic fish. A direct re- sult of this is that sexual maturity is not reached until the fish are several years old. Information on size and age at sexual maturity is set out . in Table 8. 24* Precise information is not aval] able for most species and several are estimated from the information in the lit- erature. Spawning migrations have been described in few species although there are indications generally based on negative evidence that several other species perform spawning migrations. Keysner et al. (1974) described a migration of N. rossii in May from the feeding ground north of Kerguelen to the spawning grounds to the south of the Island. The spawning grounds are characterised by high oxygen content and a rocky EOibstrate. The migration itself is thought to be in the same direction as the prevailing current which in the area has a strength of 0.6 to 1.0 kt. The post spawning northward movement in August back to the feeding grounds is thought to be made in a northerly flowing current offshore of the southward flowing current of the spawning migration. r:-28 Table 8.24, Age and Size at Sexual Maturity for Antarctic Pieh. Species Sex Sexua 1 Maturity Locality Reference Age Length (cm) Weight (e) Raja georgianus ? 60 South Georgia Permitin I969 MicromesistiuB auBtralis flii 45.3 46.4 Scotia Sea Shubnikov et al. 1969 Notothenia gibberifrons ? 35<^' 400 South Georgia Permitin and Sil'yanova 1971 Notothenia neglecta (f 8 30 Gigny Island Bverson I97O V 7 29 Signy Island cf 7 18 190 Terre Adelie Hureau I97O $ 8 22.5 300 Terre Adelie Hureau I97O N. rosBii 6' 7 48 1 700 Kerguelen Hureau I97O $ 8 55 2 700 Kerguelen N. roeeii marmorata 5(niin) 6(min) 40(min) 45(min) South Georgia Olsen 1954 N. mage 1 Ian ica 5 + 2 (6-7) 25 500 Kerguelen Hureau I97O ChampBOcephalus 4 South Georgia Olsen 1954 ChampBocephaluB (f 4 21-26 South Georgia Permitin 1973 ^ 4 21-2'3 South Georgia Chaenocephalus (9)^ South Georgia Olsen 1954 ChannichthyB rhinoceratus ^ 5 4-6 34 40-48 435 Kerguelen South Georgia Hureau I966 Permitin 1973 PseudochaenichthyB V 4-6 44-50 South Georgia Chionodraco 5 31-36 33-37 Permitin 1973 Permitin 1973 This is inferred since the authors state that only sexually mature fish were caught. In this paper mention is made only of mature fish; the size quoted is of the smallest specimen. By comparison with my own unpublished observations at Signy Island this would approximate to the size at sexual maturity. All the fish were sexually mature end aged 9 or more. 1) 2) 3) Catch rates of Chajnpsocephalus gunnari at South Georgia clearly indicate that this species moves inshore to Biwwn. Olsen (1955) failed to catch any for several months during the summer but then obtained some good hauls towards the end of March. This pattern of occ- urrence hae been confirmed in recent years by members of British Antarctic Survey fishing in Cumberland East Bay, South Georgia. A similar migration has been described for Channiohthys rhinoceratuB by Hureau (I966) although for this species the shoreward movement begins in February. G-29 Both MerlucciuB hubbBJ anrt Microni"aJ atius austral is outside the Antarctic zone apawn during the spring. Hai't (1946) showml that Morlucciue hubbai on the Patagonian Shelf moves inshore during October and November and Ciechonifjki and Weiao (1973) describe spawning in December. They also state that the eggi; cro O.O-O.9 mm diameter and are pelagic (Ciochomski and UeisB I974). The plankton survey fi'om which the above information was derived was con- ducted between latitudes ^?^ and 47 S ol F the Argentine coast and eggs were fovind throughout the region. In view of the fact that sexually mature fish have been caught in the Scotia Sea in January (Mikheyev I967) it seems highly likely that in view of the distance between these two areas spawning occurs south of 47 although there is no direct information available. Hart (1946) suggested that MicromeoistiuB aiistralis spawns 'during the spring and Weiss (1974) has confirmed this from examination of a collection of larvae. Weiss (1974) also con- siders that spawning occurs deeper than I5C m although no specific area is mentioned. The presence of mature Micromesistius austral is in the Scotia Sea during the Antarctic summer is assumed to indicate that this apocie:; migrates into the area fixsm the Patagonian region. However, although this explanation fits the observed seasonal distribution of the species in the Southwest Atlantic, it is not cotifirmed as it is possible that spawning con- centrations in the dee]) shelf zone have as y»'t gone undetected. The literature on larval stages of other Antarctic: fish is limited to a few papers des- cribing small collections (e.g., Hegan ]91C>, Hybclin 1951, LVerson I96O, Bureau I970). There is no detailed stiidy of development rates although suitable material almost certainly exists in plankton samples. 8.6 Feeding Although a detailed analysis of feeding in outside the scope of the present report, some comment should be made in order to gain a better idea of the position of fish in the ecosystem. Available information for thooe species of potential commercial importance is tabulated in Table 8.25, Tlie preponderance of Nf^totheniids, a predominantly demersal group, largely explains the high fremjenoy of benthic organisms in the diet. The presence of planktonic orga'iisms as well indicates that for some species at least there is probably some diurnal vertical feeding migration or else opportunistic feeding on planktonic organisms migrating down. lixamples of the former are probably Notothonia rossi i and Champsocephalus gunnari (see Fischer 1976) arid of the latter, Hn ja gi^orgianus and Notothenia neglecta. 8.7 Exploitation Total Catch In recent years the total reported catch within the FAO Statistical areas for the Southern Oceaji (Areas 4O, 58 ^''"^ ^^) l^^f been zero. The reason for this is the two areas knovfn to he heavily fished (South Georgia and Kerguelen) are prcsentlj' included in statisti- cal reporting areas to tlie north (4I and 51 respectively). In aridition, of the species considered in this review only two have been reported ai' separate species in the FAO Year- book of Fishery Statistics. These are Southern Pouhassou (Micromesistius nustralis) and Patagonian Hake (Merlucriuo hubbsi). There is therefore no precise information available although it has been suggested (Hureau 1973 and Ijawn unpublished Rejiort) that catches in some areas have been quite large. The following analysis is therefore based on interpreta^ tion of the reported catch in relation to the known fisheries of the area. The reported catch of Southern Poutassou in recent years is set out in Table 8.26. G-30 Table 8.25. Diot of Antarctic Fish. FiGures the porcGnta,f;;e freqi.ienc;y occurrence of a particular item. Algao X / 18 3/1 Polychaeta 10 29 / 6 3 , X Gastropoda / 1 8 Bivalvia 3 / X 19 Cephalopoda 0 '~\ / // / Amphipoda 6 10 // / 40 19 X / Isopoda 3 9 // / 51 40 Decapoda 18 3 / / 6 24 7 10 Other Crustacea 1 / / 4 1 X Dshinodermata 1 6 Other Benthos 1 / Cinderia/ 1 8 / 82 3 / 1 X Ctonophone Parathemisto h" 1.5 3 / 1 24 XX Euphausia 10 11 / 'I !0 // 1 6 II 1 3« // 48 11 // // Mysidecea ?/\ 5 / 1 8 / 7 30 ? 48 Salps X 1 / 1 Fish 16 1.5 / 3 / 38 16 / 04 3 40 15 // 30 no oO SO IG 30 so K K K 0 0 30 3G. of. :5G SG 9G 0 30 rH i-H CM m ro ^t ■J- ir> vo vo r- CO rH M -t i-i •-I (T "o to 0 (.. •H cfl' -p ■H to ■H 0 d T' to t-i (1 0 n) W J> CU 0) ^1 -t^ •H S cu si. m ,0 ■^ 0 l-l •H § I-H to -p d ^ E > •^ ro nJ rt P •r-l 0 s g •H •H 0) r-\ ■H •H 8 xt d) tn > [0 to (0

> > > ON :3 r— o O r- « 0) 0) a) o rH ON a p. C! p.' a>. j^ ;^ rH S?, ^^ rH O Q) O Q' r-~ (U 0) a a) to (-1 to f-i ^ a to -P a O ^ g 0) CO p. Li m a, ^ ^ -^ C-^li o "(J rH 0) *» to C C^' cd 1 1 c- a i-H rH (h r— LTN ,H O rH rH rH 1 1 M i-l S +J . OJ S .CI »^ ts s >! 1-1 nJ s 4^ (h rt ^ P fH .c! ri +^ 4) +5 rt P. -H Wa +^ +J +^ -^ -^ § M 4J n) 0) ^^ +» p. 0 n) rt 'd a o 0 0 0 a -H to « (•^1 0 o S Tj ,'J o o o o o O 0) a -H +5 1 1) S -P rH +^ H^ +^ +J LTN bo •H ^^ -H ■Ct S MD +^ +J +» -p O ITN 3 w .d o •HO . LOv ni ^^ "-n >o O R o O o C> OT m fn .H PI PQ m Pq >- rH o ■H « K W g m ^ Pi « -♦-> M t/5 to d CO p, CO to G -d P- rt to OT CO n} CO nJ cn to ^— « 13 t=> o >-> ti •^ !I3 D (U OJ 0) O &r « s ■rH nl •H ■d s (h rH ■^ (t! to j:^ " ^ ^ -H» d C o O o rt 3 o o a o O 01 • iS • • ^ 4) 4) [0 D-, m tn tt; tn b in en to C t3 0) s ■H '-^ d F' C .-) (3 S p — ' ir> LTN LTN 00 ^ rH "5, W J^. .H o O 13 ir^ ^■ CNI cri nl ri^ rH 1 o •^^ 1 r— ;S ^-■'^ iH iH ■& (Nl rH , — ^ ^H o ■<-t- ITN OO O • ir\ O r-l Q) Ui ^ Cvl o CM • • • rH o o O •^ ^ ^ ' — M . .-( «o| !>■ ■H t— OD rO O r^ C\J iH rO T~ o- f-l ^J- -^- irN o o ^O O rH ITN ^O \o n.t U) ^O 'O t— t— r— r— t~- t— t-- r- 0) ON ON ON ON ON C7N cr\ ON ON ON ON >H rH fH iH rH rH rH rH rH " ' " 0) •H !i C\J a. ^ w r-H C\] rH rH m •d m r-N ITN ITN ITN Ol f< +^ to nJ to 3 0) E o O ■rH ■H •H (0 ■H ," 0) ," tc P< o ^ ^< 0) (rt V rH •rt nl ■H rt O p c 4) ■H o bf o 4) Pi rH s; 4) E 21 di •a «> s-\ +^ o +» o 0) a 4) .d +^ o +> o la o ■H ■P ID O m m ^ C\J rO G-35 8.8 Biomaaa, Production and Yield The very recent build-up of fisheries in the Antarctic has meant that only limited information on biomaos and production is available and this is dinperaed in a variety of publications. Tlicre is nowhere in the literature a detailed analysis of information suit- able for fishery mana, a, o U > Ji a: H-4 INTRODUCTION 2. To encourage and stimulate investigations of the ecology and population dynamics of the organisms at different trophic levels with particular reference to krill, squid, fishes and whales. 3. To maintain liaison with FAO. 4. To advise SCAR and SCOR and through them other international organi- zations and in particular to respond to relevant recommendations of IOC and Antarctic Treaty consultative meetings. At its Executive Committee meeting in November 1975 SCOR, at the invitation of SCAR, agreed to cosponsor the group and adopted it as its Working Group 54. In order to initiate action in relation to IOC and Antarctic Treaty invitations, the first meeting of the reconstituted Group of Specialists was convened at the Scott Polar Research Institute in Cambridge, England, 6-8 October 1975 (SCAR Bulletin, 1976). During this meeting, the group agreed that its first task should be to review all existing information and to bring together knowledge of ongoing and presently planned programmes of marine biological investigations. The group also undertook the preparation of practical proposals for long term co-operative investigations of the Southern Ocean. The group welcomed the offer of the United States to host a scientific meeting on the living resources of the Southern Ocean. Accordingly, an international conference of experts was held at the US National Academy of Science Summer Studies Center in Woods Hole, Mass, 17-21 August 1976, and was followed by a meeting of the Group of Specialists (23—25 August). The chief objective of both was to review the present knowledge of the living resources of the Southern Ocean and to develop a proposal for future co-operative studies in the area. This proposal is referred to as the Biological Investigations of Marine Antarctic Systems and Stocks (BIOMASS). The research programmes included in BIOMASS are discussed in the following sections. However, before presenting these programmes, it is imperative to take into account the likely effect of marine pollution on the Antarctic ecosystem and how it relates to the proposed BIOMASS investigations. 1.1.3 Marine pollution Recently the question of exploitation of Antarctic mineral resources has come under discussion within the Antarctic Treaty Consultative Meeting and elsewhere. There is a general consensus that possible petroleum and natural gas deposits on the Antarctic continental shelf are the resources most likely to attract interest in the near future. The Eighth Antarctic Treaty Consultative Meeting in 1975 invited SCAR 'to undertake an assessment on the basis of available information of the possible impact on the environment of the Treaty area and other ecosystems dependent on the Antarctic environment if mineral exploration and/or exploitation were to occur there'. SCAR compiled a preliminary assessment as a result of various informal consultations with individual scientists and specialists and by correspondence with SCAR National Committees, based on the experience of scientific investigations in the Antarctic carried out by many nations over a number of years. This was submitted to a special preparatory meeting for the Ninth Antarctic Treaty Consultative Meeting, Paris, June/July 1976, which then invited SCAR to undertake a more detailed assessment, based on existing knowledge, for consideration at the Ninth Consultative Meeting in 1977. At its Fourteenth General Meeting in October 1976, SCAR expressed its willingness to respond to this invitation and to treat such an assessment as a task closely allied to its main objectives, although the degree of detail achievable in this further assessment would depend on the financial support available. To carry out the task SCAR established a Group of SpeciaUsts on Environmental Impact Assessment of Mineral Resource Exploration and Exploitation in the Antarctic (EAMREA), whose activities will fall into two phases: H-5 INTRODUCTION (a) the preparation of the new detailed assessment requested on the basis of existing knowledge, and the formulation of a long term research programme to fill any serious gaps in knowledge; (b) the execution of this long term research programme. It is clear that a considerable amount of research will be needed to assess the likely effects of exploration and exploitation of oil resources on Antarctic ecosvstems. While some guidance may be obtained from recent Arctic experience, there will be no substitute for research on Antarctic ecosystems and their component species. There is also a great need for studies in areas of likely activity and for basic information on the biology of key species hkely to be affected by oil pollutants. Areas of special sensitivity and/or scientific importance need to be identified and carefully planned monitoring programmes will need to be developed. From the viewpoint of the Antarctic marine ecosystem, it is clear that the BIOMASS programme wall provide much of the baseUne and other data required. There will need to be close co-ordination between other SCAR initiatives concerning the possible effects of mineral exploration and/or exploitation in the Antarctic Ocean. SELECTED REFERENCES ElSayed, S. Z. 1975. Biology of the Southern Ocean. Oceanus, Vol 18, p 40-49. SCAR Bu LLETIN. 1976. Report of a meeting of the SCAR Group of Specialists on Living Resources of the Southern Ocean. SCAR Bulletin, No 52, p 115-119. 1 .2 General objectives The need for a comprehensive international and interdisciplinary research programme in the Southern Ocean stems from the fact that the circular current system of the Southern Ocean houses a food web which is very different from those of other parts of the world ocean. Individual weights of its key members are larger by several orders of magnitude than comparable ones in other oceans, which provides the incentive for a unique mass exploitation of the Southern Ocean's key herbivore -krill. The principal objective of the BIOMASS programme is to gain a deeper understanding of the structure and dynamic functioning of the Antarctic marine ecosystem as a basis for the future management of potential living resources. Thus, we are primarily concerned with (a) contributions to man's understanding of the world ocean, and (b) developing a sound ecological strategy for the exploitation of the living resources and for the conservation of the Southern Ocean's ecosystem. To achieve these goals of basic marine science and wise ecosystem management we need to consider several objectives. The list given below emphasizes krill as the major potential food resource of the Antarctic waters and as a key element of the marine food web. At the same time, it is recognized that there are other important marine studies in the Antarctic which warrant the attention of the scientific community. BIOMASS will evolve as it proceeds and will hopefully attract new scientists with new ideas in the course of its development into a comprehensive study of the Antarctic ecosystem, its exploitation and conservation. 1. Study of the physical/chemical environment influencing krill and its food base, namely: (a) study of vertical advective and diffusive processes, as driving forces of primary production H-6 INTRODUCTION (b) Study of micro- and meso-scale horizontal processes in relation to swarming of krOl and patchiness of phytoplankton (c) study of the large scale meridional and zonal water transport at various depths in relation to the life cycle of krill. 2. Study of the variabihty between years and regions in the concentration and composition of phytoplankton. 3. Autecologjcal studies of key organisms of the Antarctic ecosystem such as whales, seals, penguins, fishes, squids, krill, some other crustaceans, salps; assessment of their standing stocks/biomass and production in selected areas of the Southern Ocean. 4. Description of the major food chains in Antarctic waters with emphasis on the flux of energy and material between the various trophic levels in selected small areas. 5. Development of models to improve our understanding of the quantitative interaction between different elements of the ecosystem and the effects of cUmate, whaling and krill fishing on structure and efficiency of the food chain. 6. Compilation and analysis of data from exploratory and commercial fishing. 7. Provision of scientific information on the Antarctic ecosystem to the scientific community, governments, industries, and other concerned bodies. H-7 2. RESEARCH PROGRAMMES 2.1. Introduction This section of the report describes the individual lines of research which are necessary for an understanding of the dynamics of the Antarctic marine ecosystem and for the conservation of the hving resources. The following sub-sections describe the proposals in respect to the main elements and processes of the system. Emphasis has been given to those elements which are believed to contribute most to the general flow of material and energy through the higher trophic levels, or are currently under exploitation or have potential for exploitation. This has inevitably meant that some elements and processes of considerable scientific interest have presently had to be neglected. So, although there is no section dealing with the benthos, this does not imply that research on benthic community structure and productivity is unimportant. Throughout the proposals, links between the benthic and pelagic communities have been emphasized. Benthic communities are important in recycling nutrients, and benthic species provide the food source for demersal fish populations. The role of benthic animals as consumers of kiill also needs to be elucidated. While an understanding of the trophodynamics of the Southern Ocean ecosystem is vital from the viewpoint of resource management and conservation, it should be stressed that we have a unique opportunity to contribute to our understanding of marine ecosystem processes as a whole. Detailed studies combined with the modelling investigations described below can lead to significant advances in our knowledge of ocean processes. Inevitably the proposals for different stocks have developed to different levels of detail. Following the long history of whaling, research on whales and the population dynamics of whales are well advanced. Over the past few years in particular, whale scientists have been actively engaged for the International Whaling Commission (IWC) and other organizations in reviewing the status of whale research, and drawing up a comprehensive international programme of whale studies. In spite of large-scale and in-depth studies of krill there are still outstanding problems. It has, therefore, been much easier to draw up a balanced and detailed programme for whales than for krill. It must also be emphasized that the krill programme (and indeed most other elements in the BIOMASS programme) will be modified in respect to specific topics as the programme develops and our knowledge of krill increases. 2.2 Modelling A whole ecosystem perspective is essential to the understanding and careful use of living resources in Antarctic waters, but it is not easy to develop this perspective from the usual starting point of observation of one particular element in the system. One method of tackling the work of understanding the whole ecosystem is the development of suitable models. They are particularly useful for ensuring a systematic approach to research and exploring the logical conclusion of certain hypotheses. Models also provide an ecologically meaningful framework for organizing, analysing, and integrating data of different types and from various sources. The power of modern computers allows highly complex situations to be handled. We already know where additional observations can be most productive. Considering such observations in an ecosystem context can provide guidance to those concerned with the management of individual resources such as whales, seals, fishes, and krill. Two levels of modelling are appropriate for study of Antarctic waters: whole ecosystem and detailed sub-models. The development of a preliminary whole ecosystem model requires H-8 RESEARCH PROGRAMMES extension of our present knowledge. Additional biological data are necessary, both field data on the biomass of ecologically important elements of the system and experimental data on transfer rates. Such a model can serve to organize existing information and to point out areas where additional research is needed. At the same time, it can be flexible enough to incorporate further information in the course of its future development. A model has been constructed of the Ross Sea ecosystem (Green, 1975) which describes the interactions of nutrients, light, ice, phytoplankton, zooplankton and larger animals. The general flow structure of this model is shown in Figure 2. For a Southern Ocean model, the main modifications needed include: (a) elements describing harvest by man, particularly of whales, krill, fish and cephalopods; (b) sub-division of certain elements (eg, of zooplankton into krill and other species); and possibly (c) division into various geographic zones, with some interchange between zones {eg, migration of whales). The Ross Sea model has been quantified, using available data and making estimates where necessary, to provide a descriptive simulation of an annual ecological cycle. ICE COMMUNITY ALGAE ICE COMMUNITY INVERTEBRATES DETRITUS AND DECOMPOSERS FIG 2. Conceptual model of the pelagic ecosystem of the Ross Sea; exchanges between the system and its environment are shown. (After K. A. Green, 1975.) Construction of one model, however complex, does not offer a complete framework from which to gain an understanding of an involved system. The development of a general model of the entire system must be accompanied by other activities, such as the development of more detailed sub-models of certain elements of the system. As an example, models which deal just with the fishes and (in a simpUfied manner) other elements interacting directly with the fishes, can simulate gross changes in fish populations. Size and age composition of the population, individual growth rates, recruitment, and variable food consumption can be included in a fish sub-model, but it would not be practicable to do this in a general ecosystem model. Similar detailed sub-models can help explain the quantitative behaviour of single components of the total system and can be used to check on the realism of the general overall model. The important sub-models on which work should start as soon as possible are: (a) the physical/chemical environment as a forcing function for phytoplankton growth, with a built-in ability to allow for different sized phytoplankton to be produced under different physical/chemical conditions; and H-9 RESEARCH PROGRAMMES (b) using phytoplankton as a forcing function, a krill model which includes the majo. predators. At present, models are most useful for perspective, data organization, and indication of information needs. When information on such aspects as the ecological role of detritus, spatial variation in productivity, or stocks of fish and cephalopods, is more complete, then predictive capability becomes an additional objective of both whole ecosystem models and sub-models. To make full use of the modelling process, it is necessary to bring the model builders together with those concerned with field studies, experimental studies, and general biological investigations. This will ensure that the models are useful and are indeed used to gain a better understanding of the system as a whole. There seems to be a need for modellers and other investigators to meet to exchange data, evaluate model results, refine model formulations for improved realism, and redefine information needs. A feedback process between model development and field studies will serve to develop the predictive capabilities of the various models as they are updated, validated and improved. It can also serve to identify information needs and provide some direction for future research. At the present state of the art of ecosystem modelling, the process of model building is as useful as model results. Objectives for Antarctic models are: 1 . Interaction among disciplines applied to Antarctic studies at both the conceptual and data analysis levels. 2. Focus for Antarctic research through organization of existing data and identification of information needs from an ecosystem perspective. 3. Descriptive ecology of the whole Southern Ocean at the level of interacting populations through dynamic simulation. 4. Feedback between model predictions and field and experimental research to refine both models and research programmes. 5. Ultimate development of a predictive model of Southern Ocean biology useful for conservation of Antarctic living resources. SELECTED REFERENCES Green, K. A. 1975. Simulation of the pelagic ecosystem of the Ross Sea, Antarctica: A time varying compartmental model. Texas A&M University, unpublished PhD thesis, 187 p. 2.3 Physical, chemical and biological environments 2.3.1 Review The broad features of the oceanography of the Southern Ocean have become well known as a result of the Discovery Investigations (1925-39) and the cruises of the Ob and the Eltanin in the late 1950's and 1960's. The knowledge accumulated from Antarctic investigations has shown that the physical/chemical settings of the Antarctic seas are unique in the world ocean. Their unique features are represented by: (a) the presence of pack ice around the Antarctic continent and the seasonal waxing and waning of the pack ice zone; (b) the variability H-10 RESEARCH PROGRAMMES of the light regime which alternates between complete perpetual darkness in winter to continuous daylight during summer; (c) the nearly uniform sea temperature which varies little with depth or season; (d) the extensive cloud cover; and (e) the circumpolarity of the surface waters and the dominant influence of the circumpolar fronts (El-Sayed and Green, 1974). It is in response to the physical/chemical environments that the Antarctic organisms have developed their characteristic features. For example, the marked seasonal variations in incoming solar radiation have a pronounced effect on plant and animal life: the rise and fall of primary produc- tion in the circum-Antarctic waters appears to be a direct response to the amount of energy received (El-Sayed, 1971). More is known about large scale surface discontinuities such as the polar front (or Antarctic Convergence) and Antarctic Divergence than about small scale discontinuities. The fact that most species appear to have a circumpolar distribution and that those south of the polar front are distinct from those to the north suggests that the biogeography of the Southern Ocean is governed largely by the dynamics of ocean circulation. The general pattern of ice cover, arrangement of water masses, distribution of currents, water column structure and nutrient salts regime are known, and it is clear from recent work that there is great variability in these parameters, both seasonally and from year to year, thaYi had previously been thought. The extent of ice cover in the Southern Ocean varies with the season and is influenced by wind; the production of cold dense bottom water may be sporadic; there is considerable variation in currents and mass transport within the West Wind Drift; a well defined clockwise eddy exists in the Weddell Sea; primary production and krill abundance vary considerably from region to region, and so on. This newly appreciated variability is currently the focus of great interest by the physical oceanographers and meteorologists participating in the ISOS programme (International Southern Ocean Studies). In the BIOMASS programme there is a need for independent research programmes in physical/chemical oceanography, and for conducting a systematic series of environmental studies on the physics, chemistry and biology of the Southern Ocean that is oriented primarily towards an understanding of the Antarctic marine ecosystem. These environmental studies will need to be closely coupled to studies of the living resources and closely co-ordinated from one ship to another and from ship to shore to satellite. Measurements by current profilers and moored current meters, as well as frequent STD casts, should provide information on vertical transport and stability and on advection. A knowledge, from sampling, of the spatial distribution and density of krill swarms in relation to the abundance and size spectra of phytoplankton, should further our understanding of the reactions of a krill swarm to its food base and the effects of grazing on the phytoplankton. Besides quantitative sampling for kriU within and outside swarms, biological observations should include controlled, quantitative plankton sampling for analyses of phytoplankton and zooplankton abundance and composition in different depth layers and at different times of day and night. Recordings of vertical distribution of chlorophyll and detritus provide additional information on the biotic environment of krill. These studies should be carried out at several selected places, mainly in different areas of high krill concentrations at the ice edge, in the neritic zone of the Antarctic Peninsula and of the Antarctic islands, as well as in the open ocean of the West Wind Drift, the East Wind Drift and the upwelling zones. If possible they should be repeated at different seasons. 2.3.2 Objectives The objectives of this section are to provide relevant information on the physical/chemical components of the marine ecosystem in the Southern Ocean as they affect the living resources and, in particular, to: H-11 RESEARCH PROGRAMMES 1. Describe the physical/chemical factors governing the distribution, abundance, productivity and behaviour of the marine organisms. 2. Describe the short and long term environmental fluctuations governing the biology of Antarctic marine flora and fauna. 3. Identify and measure, for the purpose of ecosystem modelling, the main physical and chemical driving forces in oceanic and coastal areas of the Southern Ocean. 2.3.3 Research programmes 1. Factors affecting primary production. Current estimates of primary production vary widely and our estimates are largely based on station data taken by oceanographic ships at different times and often in different years. If we are to provide reUable estimates of productivity and rates of energy flow, we must achieve geographic coverage using standardized methods and techniques. The primary production component of the BIOMASS programme will focus on a detailed documentation of the geographic variability in rates of primary productivity and on the factors which control production in the Southern Ocean. Although the productivity in polar seas with intermittent ice cover has been thought to be Umited primarily by the seasonal deficiency of available light and lack of permanent thermocUne formation, recent evidence from the Arctic/sub-Arctic regions (Apollonio, 1961; McRoy and Goering, 1974) and the Antarctic (Bunt and Lee, 1970; Meguro, 1962) indicates that the ice cover in these regions provides oceanographic conditions which in fact support high primary productivity on the under-surface of the ice during periods of the year when the water column itself is not productive. This ice mantle must, therefore, play a major role in organic production in the polar seas; and the fact that photosynthesis takes place beneath ice at low light intensities becomes of special interest. Because wind-induced turbulence carries phytoplankton down as far as the pycnocline, the amount of light available for photosynthesis varies wdth the depth of the mixed layer. Thus, a considerable amount of work will have to be devoted to the study of the factors that govern primary production in the Southern Ocean. Such study should include measurement of solar radiation, light penetration in the water column, vertical stability of the surface layer, and a suite of physical, chemical and biological parameters which influence these major variables (cloud type and cover, wind velocity and direction, transparency, organic and inorganic suspended matter, trace elements, vitamins, etc). The broad features of the phytoplankton programme are outUned below: (a) Study of the geographic and seasonal distribution of phytoplankton and estimate of primary productivity within and below the euphotic zone. Special attention needs to be given to measurements of photosynthetic activity of the micro-algae beneath the pack ice. Contributions of the various primary producer communities (pelagic, seasonal pack ice, and near -shore benthic communities— including macro-algae) should be obtained on a seasonal basis. (b) Study of the distribution and abundance of the phytoplankton in relation to physical/chemical and biological characteristics of the water masses. (c) Study of the effect of low temperature on metabolic rates of phytoplankton, and determination of their specific growth rates. (d) Study of rate Umiting processes, uptake kinetics of nitrogen and silica by organisms, and the specific role of zooplankton and in situ microbial regeneration. (e) Study of the species composition of phytoplankton. (f) Study of the grazing and dispersion loss rates (including loss rates to the benthos). H-12 RESEARCH PROGRAMMES 2. Detrital regime. There is danger in any resource-oriented research in which studies are weighted more toward the mechanisms of production than toward the equally important mechanisms of recycling the organic material produced. The main source of detritus in the open ocean is dead plankton, faecal material, and organisms embedded in the ice. It is likely that the flux of energy through these pathways is considerable. The routine detrital programme consists of measurements of particulate organic carbon which, in combination with measures of chlorophyll a (for phytobiomass) and adenosine triphosphate ATP (for living matter), allow the concentration of detritus to be determined. More complex studies include measurements of transfer rates, such as rates of release of ice detritus (by melting), rates of advection from regions of high and low detritus, rates of detrital fall out, and so on. Research on associated transfer rates, such as decomposition and mineralization, was mentioned in (1) above. 3. Factors affecting secondary production. The secondary production component of BIOMASS is focused on those elements and processes within the krill ecosystem which control or modify the flux of energy to krill. For example, part of the detrital flux may sustain significant secondary production by bacteria; other zooplankton herbivores may compete with krill for algal food; zooplankton carnivores may take developmental stages of krill in significant numbers; other zooplankton may provide alternative or supplementary forage for typical krill feeders such as some birds, seals and baleen whales. The programme should include: (a) Study of the distribution and abundance of the zooplankton species in relation to environmental gradients and to seasonal variabilities. (b) Study of the vertical distribution of the zooplankton in relation to the water masses and to the polar front. (c) Study of the role of the zooplankton in the dynamics of the pelagic food webs, including the pack ice where possible. (d) Study of the life history of key species of zooplankton with particular emphasis on their feeding biology. So far as possible, the secondary production programme will have a size specific basis, eg, size specific distribution of biomass, size specific grazing and size specific respiration rates. The role of zooplankton as forage organisms for fishes, cephalopods, birds and mammals could best be studied on board those trawlers associated with BIOMASS and at shore laboratories. Recent developments in radioactive tracer technique have made it possible to investigate food relationships within the zooplankton more effectively than in the past and such studies could well constitute a major part of research programmes on ships with well equipped biological laboratories. Because the distribution of krill corresponds, in general, with the area that is seasonally covered by pack ice, this ice-mantle is of considerable significance in the study of krill. Thus, an effective biological study of the Southern Ocean will require a fresh look at the secondary production within the pack ice ecosystem. 4. Distribution and significance of surface currents and fronts. Because of previously established macro-scale correlation between the biology of the Southern Ocean and its major frontal zones and because the distribution of krill is so patchy, the research programme on surface physical discontinuities is an important part of the BIOMASS programme. Such studies have an advantage in that they may be carried out not only by research vessels with station time at their disposal but also by supply vessels and by remote sensing from satellites. The shipboard programme mainly includes measurements of H-13 RESEARCH PROGRAMMES temperature, salinity, and water transparency with occasional nutrient analyses at interesting discontinuities. The emphasis is on continuity of observations along meridional (or near-meridional) sections. 5. Structure of the water column. The success of the krill fishery depends upon knowledge of the distribution of krill in the water column. Although empirical information is available from acoustic records, it will eventually be necessary to understand the vertical migration of krill and, in particular, the relationship between krill swarms and discontinuities in the water column. The major discontinuity at the ice edge is the shallow halocline resulting from the melting of the ice, and this should be studied. However, other discontinuities exist within the vertical range of krill, and these should also be investigated. If possible, it would be desirable to track the seasonal development and decay of these discontinuities. SELECTED REFERENCES ApolLONIO, S. 1961. The chlorophyll content of Arctic sea ice. Arctic, Vol 14, p 197-99. Bunt, J. S. and Lee, C. C. 1970. Seasonal primary production in Antarctic sea ice at McMurdo Sound in 1967. Journal of Marine Research, Vol 28, p 304-20. Ei^Sayed, S. Z. 1971. Dynamics of trophic relations in the Southern Ocean. /n: QUAM, L.ed. Research in the Antarctic. Washington, DC, American Association for the Advancement of Science, p 73-91 . Ei^SayeD, S. Z. and Green, K. A. 1974. Use of remote sensing in the study of Antarctic marine resources. In: BOCK, P., BAKER, F. W. G. and RUTTENBERG, S. eds. Proceedings of the COSPAR Symposium on Approaches to Earth Survey Problems through the Use of Space Techniques. Berlin, Akademie-Verlag, p 47-63. McROY, C. P. and GOERING, J. J. 1974. The influence on the primary productivity of the Bering Sea. /n: HOOD, D. W. and KELLEY, E. J. eds. Oceanography of the Bering Sea. Fairbanks, University of Alaska, Institute of Marine Science, p 403-21. Mf.GURO, H. 1962. Plankton ice in the Antarctic Ocean. Antarctic Record, \ol 14, p 1192-99. 2.4 KriU 2.4.1 Introduction It has been well known for many years that vast quantities of krill exist in Antarctic waters, but interest in their commercial exploitation did not arise until the early 1960's. Baleen whale stocks had declined as a consequence of overharvesting, which meant that grazing pressure on krill by these consumers was reduced. During the same period, stocks of more famiUar aquatic species, taken in traditional fishing grounds, were becoming fully exploited. Reduced opportunities for increasing harvest from other areas has focused attention on the potential of other food sources and, in particular, on Antarctic krill. Many computations have been made of production and standing stock of this pelagic euphausiid (McQuillan, 1962: Shevtsov, 1963: Jonsgard and Ruud, 1964; Mackintosh, 1966, 1970; Lyubimova and others, 1973; Omura, 1973; Fischer, 1974; Tomo and Marschoff, 1974; Everson, 1976) and most estimates of an annual harvestable yield range from 100 to 150 million tons. These figures have been derived using a variety of approaches including the consumption of past and present stocks of whales (based on their residence time in Antarctic waters, volume of stomach contents and energy requirements), consumption by other vertebrate H-14 RESEARCH PROGRAMMES predation, catch rates of krill in plankton nets and the proportion of primary production that is converted into krill. However, there are no direct or accurate data on the magnitude of either the standing stock or annual production of Euphausia superba. (Although E. superba is often considered to be synonymous with krill, and indeed is the dominant species, a number of other euphausiids should also be included under the term, and these are particularly important at the ice edge.) PLATE 1. KnW Euphausia superba. The estimated high productivity of krill has led some nations to engage m serious efforts to test its possibilities as a food source for animal and human populations. The USSR is reported to have taken some 20 000 tons in 1974, while Japan collected about 650 tons. Results of biochemical analyses and nutritional studies have encouraged commercial utilization and other nations-the Federal Republic of Germany, Poland, Norway, the United Kingdom-are also H-15 RESEARCH PROGRAMMES investigating potential krill fisheries. Recent results suggest that progress has been made towards reducing the problems of locating and catching krill, and that large catches could be taken at moderate cost, provided the fishing does not have to be interrupted to handle and process the catch. However, these latter processes still present formidable problems. Studies on a number of biological characteristics, population distribution, swarm characteristics and position of krill in the Antarctic marine ecosystem have largely been conducted during the past six decades by, among others, the United Kingdom, Norway, USSR, and Japan. While considerable information now exists concerning biological parameters, including larval stages and distribution, sexual maturity, spawning, feeding characteristics and regional differences in population densities (Bargmann, 1937, 1945; Fraser, 1937; Marr, 1962; Mackintosh, 1966, 1972, 1973; Ivanov, 1970), there is Uttle reliable direct data on krill growth rate, or growth increment and moult frequencies. Estimates concerning growth have been derived inferentially through extensive studies of class sizes, increases in mean class size with season, developmental phases (larval, adolescent, maturing and breeding adults) and their distributions with rime. The time scale from the first calyptopis to young adolescent stages seems to be established while diverse interpretations have been made from the data of post larval size frequency distribution. Mean values of krill sizes and gonadal development, derived from numerous samples collected during about 10 years and mainly between 20°Eand 70°W, have led to the conclusion that E. superba matures and breeds two summers after being spawned (Bargmann, 1945; Marr, 1962). However, samples collected in the vicinity of the island groups in the Scotia Sea in one season (February -April 1968) have been interpreted by some to demonstrate a four-year life cycle (Ivanov, 1970). With continuing efforts to determine the life span of this species. Mackintosh (1972) re- evaluated earlier data, and with additional Discovery samples reiterated previous estimates of a two-plus-year period to breeding. Thus, critical aspects of the biology and ecology of krill remain to be clarified. These include: growth rates, longevity, fecundity; hydrographic features of areas of high as well as low population densities and transport of eariy larval stages; pairing and repeat spawning. These features of krill biology must be elucidated at diverse geographic locations during every season and several annual cycles. The information derived will provide the scientific basis for judgements concerning harvestable Umits. Changes in the abundance of krill is another aspect in which full information is lacking. Though our present indirect methods have produced estimates of abundance or production which may be adequate for purposes of harvesting, the variance in these estimates (considered by some to be a factor of two or so) is so great that it is impossible to estimate the changes in the abundance of krill which have occurred or may occur. While it is populariy believed that krill stocks increased after whale stocks decreased, and there is indirect evidence that this occurred, {eg, from increases in the populations of winged birds, penguins and seals that feed on krill (Budd and Downes, 1969; Conroy, 1974)), direct evidence is lacking. Present techniques of directly estimating krill biomass, from plankton net hauls, or by acoustic methods, are subject to considerable variance because of the structure and highly irregular distribution of the swarms. Such estimates may also be biased because, for example, of the escape of larger krill from most nets, the fully unknown proportion of krill outside swarms and the lack of information on the acoustic properties of krill, either individually or in swarms. Though these problems are of some significance, they do not invalidate present estimates of biomass and productivity which have been presented as broad ranges. They do make it difficult or impossible, however, to achieve the precision needed to monitor changes in the krill population which might occur as a result either of a recovery of the whale stocks or of large scale exploitation. The development of methods of monitoring changes in krill abundance is highly desirable. Techniques that are under investigation and at the present stage show some promise include refined acoustic methods from ships or buoys, aerial surveys and the use of sateUites. H-16 RESEARCH PROGRAMMES 2.4.2 Objectives Because of its key position in the Antarctic ecosystem and the possibility of krill making a significant contribution to world food supply at some time in the future, the studies of krill proposed here have a significant part in the whole BIOMASS programme. It is planned that these studies will lead to a better understanding of: (a) the fundamental biology off. superba; (b) the extent of dependence of higher trophic levels on krill stocks; (c) the interrelations between krill swarming and the hydrographic features of Antarctic waters. The studies have been drawn up so that in addition to improving our general store of scientific knowledge, sufficient information will be available before any intensive exploitation of krill begins. If these objectives are successful it should be possible to make use of the large krill resource without risk of lasting damage to stocks, or to stocks of mammals, birds and other animals which depend to a greater or lesser extent on krill for food. In particular the objectives of the programme will be to: 1 . Make an intensive study of the life history and population dynamics of krill. 2. Elucidate the contribution of krill to the dynamics of its main predators. 3. Develop systems to monitor changes in the abundance and composition of the krill stocks. 2.4.3 Research programmes It would not be possible, given the scales of the problems and the magnitude of the resources that are likely to become available for the BIOMASS programme, to draw up practicable proposals that would deal with all the objectives of the krill research programme in the near future. The present proposals, therefore, have been selected to deal with topics that are particularly important in themselves or lay the groundwork for later important investigations. The specific proposals are: 1 . Determine growth rate and longevity using (a) laboratory and field experiments, and (b) collection and analysis of length composition data for krill collected in all seasons and areas. 2. Describe the spatial distribution of krill with the objectives of determining ranges of biomass densities and the shape, extent and stability of swarms, especially diel changes. 3. Develop acoustic techniques for the quantitative survey and evaluation of krill stock. 4. Carry out surveys with quantitative nets (RMT, Bongo, etc) to determine the vertical and horizontal distriburion of krill in oceanic areas, coastal areas and the pack ice zone. 5. Determine the types of fishing effort data {eg, number of hauls, searching time) from which a catch per unit effort could be produced to provide the most valid index of abundance, and ensure that such data are collected from any commercial krill fishing operations. 6. Carry out ecological studies throughout the Southern Ocean, with particular emphasis on the Weddell sector, in the most likely areas of potential intensive exploitation and in the important but poorly understood area of the East Weddell Drift. 7. Identify stocks or racial differences in all study areas. If sub-groups are discovered, conservation measures and management of commercial harvesting will be expected to reflect growth characteristics of such groups. 8. Study the distributions, vertical and horizontal transport of early larvae Teees to first calyptopis stage). H-17 RESEARCH PROGRAMMES PLATE 2. A 25-ton catch of krill being hauled onto the deck of FFS Walther Henvig. H-18 RESEARCH PROGRAMMES 9. Investigate the biochemical composition of krill and its relation to feeding, season and growth. 10. Investigate the spawning behaviour, area, depth and intensity, and the possible occurrence of repeated spawning. 1 1 . Evaluate the possibility of using remote-sensing techniques for monitoring krill. In addition it will be important to ensure that comprehensive catch and effort statistics are collected and reported to the appropriate international agency. The handling of practical fishery information can be considered as the responsibility of fishery institutes and agencies. The information is often needed to achieve the best productive tactics for the development and utilization of kriD, but if large exploitation starts it will also be invaluable for quantitative studies of krill dynamics. During the Woods Hole conference, it was possible to examine acoustic techniques in some detail. While the commercial fishing expeditions have clearly demonstrated the efficiency of hydro-acoustic techniques in detecting swarms of krill, the translation of such signals into absolute units requires special study. It was proposed to conduct as soon as possible a small scale study for the purpose of: (a) determining the back scattering strength of various densities of krill by cage experiments (b) providing a calibration coefficient from simultaneous fishing and hydro-acoustic mapping (c) determining the variance encountered in different survey strategies. To implement these proposals close international co-operation will be needed. Useful co-operation has already developed during recent years between scientists of several countries engaged in krill research and related aspects; it ranges from exchange of information such as cruise plans, prehminary results and publications, to joint participation on board research vessels during krill surveys. The existing international collaboration in krill research should be further expanded and intensified. It is essential that the methods of sampling, recording and reporting be standardized or intercalibrated as far as possible, in order to avoid the collection of material which might turn out later to be incomparable. Working groups on methods of krill research should be established to consider all items requiring standardization (eg, length measurements, maturity stages, stomach filling), and to elaborate proposals for survey techniques, data collection, evaluation and reporting. The recent establishment of SCOR Working Group 52 on the estimation of micronekton abundance is a first step in this direction. SELECTED REFERENCES BaRGMANN, H. E. 1937. The reproductive system oi Euphausia superba. Discovery Reports, Vol 14, p 325-50. BaRGMANN, H. E. 1945. The development and life history of adolescent and adult krill, Euphausia superba. Discovery Reports, Vol 23, p 103-78. BUDD. G. M. and DOWNES, M. C. 1969. Population increase and breeding in the Kerguelen lur Seal, Arctocephalus Iropicalis Gazella, at Heard Island. Mammalia, Vol 33, p 58-67. CONROY, J. W. H. 1975. Recent increases in penguin populations in Antarctica and the sub-Antarctic. In: STONEHOUSE. B. ed. The biology of penguins. London. Macmillan, p 321-36. EVERSON, I. 1976. Antarctic krill: a reappraisal of its distribution. Polar Record, Vol 18, No 112, p 15-23. FiSCHKR, W. 1974. Der Kiill (Euphausia superba) and andere Nahrungsreserven im Gebiet der Antarktis (Krill (Euphausia superba) and other food resources in the Antarctic area]. Protokolle zur Fischereitcchnik, Bd 62, p 226-88 (translated). H-19 RESEARCH PROGRAMMES FraSER, F. C. 1937. On the development and distribution of the young stages of krill {Euphausia superba). Discovery Reports, Vol 14, p 1-92. GULLAND, J. A. 1970. The development of the resources of the Antarctic seas. In: HOLDGATE, M. W. ed. Antarctic ecology. Vol 1. London and New York, Academic Press, p 217-24. GuLLAND, J. A. 1976. Antarctic baleen whales: history and prospects. Polar Record, Vol 18, No 112, p5-13. HeMPEL, G. 1968. Area reviews on living resources of the world's oceans. Antarctica. FAO Fisheries Circjjlar, No 109.3. IVANOV, B. G. 1970. On the biology of the Antarctic krill, Euphausia superba. Marine Biology, Vol 7, p 340-51. Johnstone, G. W. and .Murray, M. D. 1972. Dominican gulls in the Australian Antarctic Territory. Australian Bird Bander, Vol 10, p 59-60. JONSGARD, A. and RUUD, J. T. 1964. Studies on the southern stocks of Blue and Fin whales. In: CARRICK, R., HOLDGATE, M. W., and PREVOST, J. eds. Biologic Antarctique. Paris, Hermann, p 333-39. Lyubimova, T. G. and others. 1973. Prospects of the utilization of krill and other non-conventional resources of the world ocean. By T. G. Lyubimova, A. G. Naumov, and L. L. Lagunov. Fisheries Research Board of Canada. Journal, Vol 30, No 12, Part 2, p 2196-2201. Mackintosh, N. A. 1966. The swarming of krill and problems of estimating standing stock. The Norwegian Whaling Gazetteer, Nr 11, p 213-16. Mackintosh, N. A. 1970. Whales and krill in the twentieth century. In: HOLDGATE, M. W. ed. Antarctic ecology, Vol 1. London and New York. Academic Press, p 195-216. Mackintosh, N. A. 1972. Life cycle of Antarctic krill in relation to ice and water conditons. Discovery Reports, \o\ 36, p 1-94. Mackintosh, N. A. 1973. Distribution of post-larval krill in the Antarctic. Discovery Reports, Vol 36, p 95-156. MaRR, J. S. W. 1962. The natural history and geography of Antarctic krill {Euphausia superba Dana). Discovery Reports, Vol 32, p 33-464. Mcquillan, H. 1962. The Antarctic krill. Western Fisheries, Vol 63, No 4. Moiseyev, p. a. 1970. Some aspects of the commercial use of the krill resources of the Antarctic seas. In: HOLDGATE, M. W. ed. Antarctic ecology, Vol 1. London and New York, Academic Press, p 213-15. Moiseyev, P. A. 1971. Development of scientific bases of fishery and of the methods controlling the processes of biological productivity in the oceans. In: Fundamental biological productivity in the oceans and its exploitation. Washington DC, National Marine Fisheries Service, Foreign Fisheries Division, p 7-11. (Translation No 72-50071A.) NeMOTO, T. 1968. Feeding of baleen whales and krill, and the value of krill as a marine resource in the Antarctic. In: CURRIE, R. I. ed. Symposium on Antarctic Oceanography. Cambridge, Scientific Committee on Antarctic Research. OmURA, H. 1973. A review of pelagic whaling operations in the Antarctic based on the effort and catch data in 10° squares of latitude and longitude. Research on Whales Institute. Scientific report, p 105-21. ShEVTSOV, V. V. 1963. Certain data on Antarctic kriU. UNIRO, No 12, p 22-33 (translated). SlADEN,W. J. L. 1964. The distribution of Adelie and Chinstrap pengums. In: CARRICK, R., HOLDGATE, M. W., and PREVOST, J. eds. Biologic Antarctique. Paris, Hermann, p 359-65. TOMO, A. and MaRSCHOKF, E. 1974. Estimation of those parameters of the population of E. superba which would be important for the management of this resource. Buenos Aires, Instituto Antartico Argentine. (Contribucion N:200.) ZeNKOVICH, B. a. 1970. Whales and plankton in Antarctic waters. In: HOLDGATE, M. W. ed. Antarctic ecology. Vol I. London and New York, Academic Press, p 183-85. 2.5 Marine mammals and birds 2.5.1 Introduction The two groups of marine mammals to be found in the Southern Ocean are the seals and whales. Both represent substantial potential sources of food: the seal stocks are, and the whales were vastly more important than their Northern Hemisphere counterparts in terms of number and biomass. While there are no autochthonous peoples engaged in hunting in the south as there H-20 RESEARCH PROGRAMMES are in tlie north, it is well known that the stocks of large whales have been commercially overfished in the Southern Ocean since 1904, and with the exception of the Minke Whale are still very low compared with their former abundance. There is a need for proper conservation and management and a growing awareness of the complex interactions between seals and whales involving possible competition for food with each other and with birds, fishes and squids. Two international conventions are concerned with the marine mammals: the International Whaling Convention, dating from 1931, and the Convention for the Conservation of Antarctic Seals, signed in 1972 but not yet in force. The SCAR Group of Specialists on Seals and the Subcommittee on Bird Biology of the SCAR Working Group on Biology are also active in this field. Adequate conservation and management depends on a continuing input of data and information. 2.5.2 Ecology and abundance 1. Seals The pack ice region around Antarctica is the habitat of the Antarctic seals. These are the Crabeater Seal Lobodon carcinophagus, the Weddell Seal Leptonychotes weddelli, the Leopard Seal Hydrurga leptonyx. and the Ross Seal Omatophoca rossi. These species have recently drawn considerable worldwide attention since they are a resource as yet unexploited by man. Other seal species which inhabit the Southern Ocean are the Southern Elephant SQzXMirounga leonina and the Fur Seals Arctocephalus gazella and A. trapicalis. ihese species were overexploited by the 18th and 19th century sealers and the stocks are still recovering. Table 1 . Crude estimates of antarctic seal populations, biomass and food consumption Species Stock (thousands) Mean wei (kg) ght Population biomass' Annual food total krill consumption' squids fishes Elephant 600 500 300 6 000 — 3 000^ 3 000 Leopard 220 272 60 1 403 519 112 182 WeddeU 730 246 180 4 211 - 463 2 232 Crabeater 15 000 193 3 000 67 245 63 210 I 345 2 017 Ross 220 173 38 892 80 571 196 Fur 350 50 17.5 410 205 102 102 TOTAL^ 17 000 _ 3 600 80 000 64 000 6 000 8 000 Thousands of metric tons. Assumed proportions in absence of pelagic data. ^Totals rounded. Scientific studies on Antarctic seals began in tlie early 1900's. Most of the work since has been carried out from icebreaker support vessels or in close association with the Antarctic shore stations. In general, the Crabeater, Leopard and Ross seals are distributed in the pack ice regions surrounding the Antarctic continent. Erickson and Hofman (1974) have provided maps which summarize information on distribution and abundance of these species based on census information from the years 1968 and 1974. Estimates of the abundance of various species in these regions are presented in Table 1. For the pack ice species the estimates have been corrected for variation which might occur in surface counts due to the seals' 24-hour activity pattern, but they are, nevertheless, underestimates because an unknown proportion is in the water even at peak haul out times. H-21 RESEARCH PROGRAMMES (a) Crabeater Seal. The Crabeater Seal is the most abundant seal in the Antarctic and probably in the world. Siniff and others (1970) and Erickson and others (1971) have shown a relationship between ice type, in terms of cover and density, and Crabeater Seal abundance. The largest numbers of Crabeater Seals are to be found in small ice floe environments covering between 30 and 70 per cent of the water surface. The Crabeater Seal has a 24-hour activity pattern with peak numbers hauled out, it is thought, at local apparent noon; research is needed to document what the proportion may be. The food of the Crabeater Seal is mainly krill, mostly Euphausia superba, though there is a possibility that the bulk of the food will be E. crystallorophias in some areas over the shelf. PLATE 3. A Crabeater Seal family group. Little is known about the breeding of the Crabeater Seal since it occurs in the austral spring when the ice is at its maximum (Laws, 1958; 0ritsland, 1970). After weaning, the pups probably disperse and form groups in the pack ice. Hofman (1975), Lindsey (1937) and Bertram (1940) suggest that the young of the year congregate along the edge of the Antarctic continent in areas where the ice is quite dense. This leads to a segregation by sex and age, the older, mature animals being at the periphery of the pack ice with the younger animals toward the interior (Hofman, 1975). The maximum recorded age for a Crabeater Seal is 33 years, little is known about mortahty, although the Killer Whale and Leopard Seal are known to feed on this species. Predation would appear to be significant since a high proportion of Crabeater Seals observed in the pack ice are scarred. (b) Leopard Seal. The Leopard Seal is the largest of the true Antarctic seals, it appears that it is catholic in its H-22 RESEARCH PROGRAMMES diet and takes penguins under rather specific conditions but utilizes kriU, fish and other seals, particularly young Crabeater Seals, when conditions are favourable. Little is known about the life cycle of the Leopard Seal. Pupping and mating probably occur in the pack ice. Oritsland (1970) reported that males and females become sexually mature between three and six years with the maximum hfe span in the vicinity of 25 years. (c) Ross Seal. Information on the Ross Seal, known as the rarest and least studied of the Antarctic seals, has been summarized by Hofman and others (1973). It appears mainly to inhabit pack ice of 5/10-7/10 cover. Most sightings have been of solitary individuals but groups of up to 13 have been sighted (Laws, 1964). The Ross Seal is extremely rare in the Weddell Sea but its distribution is uneven: much higher concentrations are found in the Amundsen, Bellingshausen and King Haakon VII seas. It is considered to account for between one and two per cent of the total Antarctic pinniped population. Cephalopods have been reported to be the main food of the Ross Seal although both fish and krill have been found in its stomachs. Information on reproduction is limited, most coming from Oritsland (1970) who examined 15 specimens. It appears that pupping and mating occur in November and December. (d) Weddell Seal. The Weddell Seal is the best known of the Antarctic seal species as it is found near Antarctic shore stations where access for study has been possible. It aggregates during the pupping season along perennial tide cracks where pups are born on the surface of the ice. Pupping occurs as early as September in northern areas and in late October and early November in the more southern areas. Weaning is usually accomplished by the sixth or seventh week, at which time the colony disperses. The males occupy underwater territories (D. B. Siniff, unpublished ms) along the tide crack; the exact time of mating is uncertain but it appears to be near weaning. As with the other Antarctic seals, peak haul out is around local apparent noon. Their food includes fish, crustaceans (including euphausiids), and cephalopods (Dearborn, 1965). At McMurdo Station the Weddell Seal feeds on fish of the genus Trematomus and midwater species such as Pleuragramma antarcticum. (e) Fur Seals. Although formerly hunted to near extinction, there is at present no harvest of these species and the populations are increasing. The Fur Seal that breeds on islands south of the Antarctic Convergence is Arctocephalus gazella (Repenning and others, 1971). The main population oi A. gazella is found on South Georgia. It has increased from at most a few hundred in the 1930's to around 350 000 now. Subsidiary smaller populations are found on other islands of the Scotia Arc north of 65°S, Bouvet and McDonald islands and, lately, this species has also begun breeding at Marion Island, sympatrically with the more northern form>4. tropicalis (Bonner, 1968; Erickson and Hofman, 1974). The species may be expected to reach its former abundance, measured in millions, in 15—20 years time. A. tropicalis, the Fur Seal of the islands to the north of the Antarctic Convergence, has shown a slower rate of recovery. Stocks at Marion Island have increased from around 500 to 7 000 between 1950 and the present, and at Gough Island from about 1 2 000 to 26 000 in the same period (P. R. Condy, personal communication). Fur Seals come ashore to breed on rocky coasts in November and lactating females remain in the vicinity of the rookeries until the following March. For A. gazella food at this period appears to be mainly Euphausia superba (though fishes and squids are also taken). The food of A. tropicalis is less well known but includes fishes, squids or the rock lobster Jasus sp. After H-23 RESEARCH PROGRAMMES harem break-up the Fur Seals disperse. Their distribution and feeding habits in the winter are unknown. (0 Southern Elephant Seal. This species is similar to the Southern Fur Seal in that it congregates on beaches to have pups and mate. The total mid-year population has been estimated at about 600 000 with the largest population (370 000) on South Georgia (Laws, 1960). The first mature bulls establish territories in September and the breeding season lasts until mid-November. Elephant Seals do not feed during the breeding season nor during the period of moult that follows. Their primary food is cephalopods and fish, but in what proportion is not known. Although almost exterminated at South Georgia in the 19th century, the numbers under protection were increased and the species was the subject of a successful nationally managed industry, which took only adult males from 1910 to 1964; the sustainable yield appeared to be about 6 000 (Laws, 1960). PLATE 4. Minkc Whales in Normanna Strait, South Orkney Islands. 2. Antarctic whales. Tlie whales include the baleen whales (Mysticeti) and toothed whales (Odontoceti). The waters surrounding the Antarctic continent are inhabited by the following five species of the large baleen whales: Blue Whale Balaenoptera musculus. Fin Whale B. physalus. Sei Whale 8. borealis. Humpback Whale Megaptera iiovaeangliae and the Sperm S^\vd\e Fhyseter catttdon. the only large odontocete. The smaller Minke Whale Balaenoptera aaitorostrata also inhabits these waters, and the Southern Right Whale Eubalaena australis occasionally migrates into the Z o So. u (0 H-24 RESEARCH PROGRAMMES E 3 2 ■2° I 00 ^ oor-«rjooo mr^sD o H o c < " < o A. OC^ — OOO "O — 00(SO JO _ C00»rt000 .. -,._^ — C^ b CN CO 00 (N O O 5 o (I. Q OV ,-'^a oor-oooo r^joo^a-oo COH fflS^cS OOOOOOOO ror*)or-00 H Q SS o W < So „^ 3 3 j: S •o u 0 x> w {J (J C A c 00 c > < 4J c J= o t u c ^^ u « ■o xn o 4) sS •O Uea OOv^OOv^vi o M c £ s £ g o E o V a X T3 CO E ii o S t; c J= £ s f ^ S a V < c 1> Blue Sei Hum Mink H a n [1. O 03 C/5 b S £ S -^ d S " ^ -- m en - n n Biomass (thousands) Spheniscidae 121 770 520.7 Diomedeidae 12 630 44.45 Procellariidae 39 354 10.17 Pelecanoididae 1 1 000 1.3 Hydrobatidae Phalcrocoracidae 2 835 0.56 Stercoraciidae l.aridae TOTAL' 188 000 580 H-28 RESEARCH PROGRAMMES colonial birds, the major limiting factor may be availability of nest sites, there is evidence indicating that food availability is still an important factor. In terms of food consumption, birds and large whales are each estimated to take about 40 million tons annually while seals take about twice as much--80 million tons. OriginaUy the large whales were estimated to have taken nearly 200 million tons annually of krill, squid and fish before their stocks were depleted by whaling. The staggered southerly migration of baleen whales and their feeding preferences for different class sizes of krill and other zooplankton reduce competition. However, it has been demonstrated that longitudinal segregation also occurs with the larger and older whales arriving first to occupy certain areas of the feeding grounds, and younger, later arrivals being displaced to the periphery (Laws, 1960). At South Georgia, for example, where the different species overlap, competition for krill of the same size occurs. The growth and pregnancy rates of Blue and Fin whales have increased and the mean age of sexual maturity has decreased during the periods of expanding whaling operations in response to increased food abundance (Laws, 1977 and in press). Similar increases in pregnancy rates and advancement of maturity in Sei Whales have been shown to precede large scale exploitation of the species, and so could not be a direct result of hunting. This lends support to the belief that the increased early pregnancies and growth rates are indirect responses to whaling through its effect on food supply. There is no direct evidence of a change in abundance of krill-eating Crabeater Seals, but there is evidence of an increased rate of reproduction. Their mean age of sexual maturity in Whaling Zone I (the Sanctuary) has decreased from 4 years in 1955 to 2.5 in 1970 during a period of rapid depletion of whale stocks (Laws, 1977). At South Georgia the population explosion shown by the krill-feeding Fur Seal Arctocephalus gazella is very marked and other colonies are developing in the Scotia Arc. Their distribution overlaps the baleen whale feeding distribution and the much more rapid recovery of -4. gazella in comparison with y4. tropicalis can probably be attributed to changes in krill availability. 2.5.4 Objectives. The objectives of the programme will be: 1. Identification of unit breeding or management stocks, their location, movements and possible mixing. 2. To obtain full data by species, sex, age and size for all catches. 3. Stock assessment and the estimation of potential yields when appropriate. 4. An understanding of trophodynamics. 5. An understanding of interactions between whales, seals and birds and with other groups at a similar trophic level, such as fishes and squids. 6. Formulation of suitable management and conservation procedures. Objective 2 is being met under schemes operated by the Bureau of Whaling Statistics, and in respect to seals and birds by the Antarctic Treaty (Agreed Measures for the Conservation of Antarctic Flora and Fauna). The Convention for the Conservation of Antarctic Seals calls for reporting of catches. 2.5.5 Research programmes 1. The mapping of seasonal pack ice distribution and type is particularly important in extrapolating sample seal densities to larger areas and, again primarily for seals, in assessing the significance of residual pack ice regions in summer for the separation of stocks. (Objectives 1 , 2 and 4) 2. Research should be carried out throughout the range of species including, for whales and birds, the winter period outside the Southern Ocean. H-29 RESEARCH PROGRAMMES 3. Observations should be made from ships, including icebreakers, and aircraft, to establish distributions, estimate densities and absolute abundance of, in particular, protected whale species such as the Blue and Humpback, seals and birds in the pack ice, and birds and whales in the open sea. Seals cannot be counted in the water and counts on the pack ice give a minimal estimate of abundance. To resolve this problem, behavioural observations in the pack ice will be necessary using the telemetry developed for studies of Weddell Seals in fast ice areas. If the age structure of the population is known, estimates of the abundance of birds, Elephant, Weddell or Fur seals, a portion of whose stocks concentrates on land or fast ice to breed, can be made by extrapolation from counts of breeding birds or seal pups. These counts can be made on the ground or from aerial photographs, but must be carried out before the breeding birds or seal pups disperse. Differential dispersal at sea by season, sex and age means that direct counts of the total population of land breeders are not possible. Information on the distribution and abundance of small whales is almost totally lacking and indirect methods such as radio- and sonic-tagging and hydrophone arrays must be employed. (Objectives 1 and 3) 4. Information from scientific research needs to be supplemented by data from whaling and sealing operations. The most direct way of evaluating estimates of sustainable yields is by using data on catches and effort to determine the effect of various levels of exploitation on the stocks. A programme of pelagic research captures from specially equipped whaling vessels such as the Norwegian Peder Huse, which was a combined whale catcher and factory ship, and from other vessels, including icebreakers for the seals in the pack ice, is necessary to complement and extend the results from commercial operations. (Objective 2) 5. Existing marking programmes should be continued and new ones developed, especially for smaller species. Methods should also be developed for the study of individual whales, seals and birds, possibly including satellite tracking, radiotelemetry and sonic tagging. (Objectives 1 and 3) 6. Marking/recapture data, studies of population genetics, morphometries and other indicators of discrete populations should be used to establish unit stocks. Much can also be learned from captured seals without having to kill them-for example, by collection of blood samples for serological and reproductive studies, and of nails for age determination. (Objectives 1 , 2 and 4) 7. For animals killed during commercial operations, in research operations using special ships and in shore-based work, the following basic studies are required: (a) Morphometries. (b) Age determination to establish growth rates, population structure and natural mortality rates, and age-specific reproductive parameters. Good age criteria exist for a number of species from earlier studies; for example, annual layers in teeth (dentine or cement), nails or ear plugs. Because of the numerous age groups, relatively large samples will be needed. (c) Analysis of reproductive state and reproductive history. This would include back-calculation of age at puberty of marine mammals from ear plug and tooth structure to establish possible trends in the mean age at puberty. Age-specific and time-specific pregnancy rates are also required for the mammals and, similarly, breeding success for birds. (d) Investigation of food habits and ecological separation by species, sex and age. Preliminary studies indicate a more or less clear ecological separation in terms of geographical distribution, feeding depth stratification, adaptations for feeding (morphology, visual pigments) and food preferences. More detailed quantitative information is needed to refine existing knowledge and to establish possible seasonal and geographical differences. H-30 RESEARCH PROGRAMMES (e) The development of programmes for estimating metabolic rates and energy flow. Just as biomass is a better measure than numbers for estabUshing the significance of a species in the ecosystem, so energetic terms are preferable to biomass, particularly in studies of interactions and competition for a possibly limiting food resource. (Objectives 1,3,4 and 5) 8. Analysis of the various parameters (stock sizes, growth, mortality, reproductive rates, feeding patterns) should be continued in an attempt to discover how they vary with changes in the abundance of seals, whales, birds and other groups. This will not, however, lead to early results. (Objectives 5, 6) 9. Finally, better and more dynamic ecological and trophodynamic models and population models are needed to help in the formulation of conservation and management strategies. (Objectives 4, 5 and 6) SELECTED REFERENCES Barratt, a. and MoUGIN, J. L. 1974. Donnees numeriques sur la zoogeographie de I'avifaune antarctique et subantarctique. CNFRA (Paris), No 33, p 1 -18. Bertram, G. C. L. 1940. The biology of the Weddell and Crabeater sels, with a study of the comparative behaviour of the Pinnipedia. British Graham iMnd Expedition, 1934-37, Scientific Reports, Voll, No 1, 139 p. (London, British Museum.) Bonner, W. N. 1968. The Fur Seal of South Georgia. British Antarctic Survey Scientific Reports, No 56, 81 p. Bonner, W. N. and Laws, R. W. 1964. Seals and seaUng./n: PRIESTLEY, R., ADIE, R. J., a«(/ ROBIN, G. de Q. eds. Antarctic research. London, Butterworth, p 163-90. British Antarctic Survey. 1977. British Antarctic Survey, internal Report. (Unpublished.) Brownell, R. L. 1974. Small odontocetes of the Antarctic. Antarctic Map Folio Series, Folio 18, p 13-19. CaRRICK, R and IngHAM, S. E. 1967. Antarctic sea-biids as subjects for ecological research. In: NAGATA, T. ed. Proceedings of the Symposium on Pacific-Antarctic Sciences . . . Tokyo . . . 1966. Tokyo, National Science Museum, Department of Polar Research, p 151-84. Dearborn, J. H. 1965. Food of Weddell Seals at McMurdo Sound, Antmcticdi. Journal of Mammalogy, Vol 46, No l.p 37-43. Erickson, a. W. and others. 1971. Distributional ecology of Antarctic seals. By A. W. Erickson, D. B. Siniff, D. R. Cline, and R.J. Hofman. In: DEACON, G. ed. Symposium on Antarctic Ice and Water Masses. Cambridge, Scientific Committee on Antarctic Research. Erickson, A. W. and Hofman, R. J. 1974. Antarctic seals. Antarctic Map Folio Series, Folio 18, p4-13. Gambell, R. 1976. World whale stocks. Mammal Review, Vol 6, No 1, p 41-53. Hofman, R. J. 1975. Distribution patterns and population structure of Antarctic seals. University of Minnesota, PhD thesis. Hofman, R. J. and others. 1973. The Ross Seal (Ommatophoca rossi). International Union for the Conservation of Nature. Research Publications (New Series), No 39, p 129-39. International Whaling Commission. 1950-1974. Report of the International Whaling Com- mission, Nos 1 -24. Laws, R. M. 1958. Growth rates and ages of Crabeater Seals, Lobodon carcinophagus. Proceedings of the Zoological Society of London, Vol 1 30, Part 2, p 275-88. Laws, R. M. i960. The Southern Elephant Seal {Mirounga leonina (Linn.)) at South Georgia. Norsk Hvalfangst-Tidende, Arg 49, Nr 10, p 466-76, Nr 11, p 520-42. Laws, R. M. 1964. Comparative biology of Antarctic seals. In: CARRICK, R., HOLDGATE, M. \J.,and PREVOST, J. eds. Biologie Antarctique. Paris, Hermann, p 445-54. Laws, R. M. 1977. Seals and whales of the Southern Ocean. In: FUCHS, V. E. and LAWS, R. M. eds. Scientific research in Antarctica. Philosophical Transactions of the Royal Society (London), B, Vol 279, No 968, p 81-96. Laws, R. M., in press. The significance of vertebrates in the Antarctic marine ecosystem. In: LLANO, G. A. ed. Adaptations within Antarctic ecosystems: Proceedings of the 3rd SCAR Symposium on Antarctic Biology. Houston, Gulf Pubhshing Company. H-31 RESEARCH PROGRAMMES LiNDSEY, A. A. 1937. The Weddell Seal in the Bay of Whales, Antarctica. Journal of Mammalogy, Vol 18, p 127-44. Mackintosh, N. A. 1965. The stocks of whales. London, Fishing News (Books) Ltd. OrITSLAND, T. 1970. Seahng and seal research in the south-west Atlantic pack ice, Sept. -Oct. 1964. /«: HOLDGATE, M. W. ed. Antarctic ecology. Vol 1. London and New York, Academic Press, p 367-76. RepENNING, C. a and others. 1971. Contribution to the systematics of the Southern Fur Seals, with particular reference to the Juan Fernandez and Guadalupe species. By C. A. Repenning, R. S. Peterson, and C. L Hubbs. In: BURT, W. H. ed. Antarctic Pinnipedia. Antarctic Research Series. Vol 18. SiNIFF, D. B.and others. 1970. Population densities of seals in the Weddell Sea, Antarctica, in 1968. By D. B. Siniff, D. R. CUne, and A. W. Erickson. In: HOLDGATE, M. W. ed. Antarctic ecology. Vol 1. London and New York, Academic Press, p 377-94. Watson, G. E. and others. 1971. Birdsof the Antarctic and sub-Antarctic. By G. E. Watson, G. E. Angle, J. PhiUip, P. C. Harper, M. A. Bridge, R. P. Schlatter, W.L.N. TickeU, J. C. Boyd, and M. M. Boyd. Antarctic Map Folio Series, FoUo 14. 2.6 Fishes 2.6.1 Introduction A large number of species of fish occur in the Antarctic, but most of these are small or few in numbers (Andriyashev, 1965; Dewitt, 1971; Everson, 1970; Hureau, 1970). At the present level of our understanding, Antarctic fishes do not significantly contribute to the general dynamics of the Antarctic ecosystem. Attention, therefore, has been concentrated on the following 12 species which are believed to be of major importance in the Southern Ocean (see also Figures 3, 4 and 5). (a) Notothenia rossiL Found in all coastal areas of the Scotia Arc, from South Georgia south to the Antarctic Peninsula; at Kerguelen, Crozet, Macquarie and Marion islands, and probably also at Bouvet Island. It grows to about one metre in length. The species at South Georgia spawns during April or May, and at Kerguelen at the end of May. It has probably been the most heavily exploited of Antarctic fishes. However, since the FAO statistical reporting areas combine South Georgia and the Kerguelen islands with the respective areas to the north, and because A', rossii is not on the list of reported species, it is combined with the 'other demersal percomorphs' category. Estimates of catches at South Georgia range from 400 tons to something in excess of 400 000 tons annually. (b) Champsoceptialus gunnari. Distribution essentially the same as for A', rossii. The species grows to about 60 to 70 cm. Information about spawning is not available. Recent expeditions by the Federal Republic of Germany and Poland have found commercial quantities. It is one of the major species believed to have made up catches totalling 120 000 tons taken around the Kerguelen archipelago during 1971-72. (c) Micromesistius australis. Found in the Magellanic and New Zealand regions and known to migrate south into the Antarctic Zone during the summer to feed on krill. It reaches a length of over half a metre and spawns north of the Antarctic Zone during the spring. Total annual catches up to 7 900 tons have been reported for the south-west Atlantic Ocean, which probably represent fishing efforts in the Scotia Sea. (d) Notothenia gibberifrons. Found throughout the Scotia Arc region and off the Antarctic Peninsula. It reaches a length of about half a metre. Information about spawning is not available. Also not available are catch statistics, but the recent West German expedition took large numbers, and circumstantial evidence indicate^ *hat the USSR may be using it for at least local consumption. H-32 RESEARCH PROGRAMMES HotothanLa loa-ii (length to 1 m) Champsoaephalus gunnari (length, to 60-70 cm) Micromesistius austfalis (length to SO cm) Notothenia gibberifrons (length to 60 cm) FIG 3. Antarctic fishes in the order discussed in the text. H-33 RESEARCH PROGRAMMES (e) Pseudochaenichthys georgianus. Found at South Georgia, the South Orkney Islands and off the Antarctic Peninsula. It reaches a size of about half a metre or a little more. No information about spawning or exploitation is available except that recent West German and Polish expeditions caught significant quantities. (f) Dissostichus mawsonL Circum-Antarctic, both in the open ocean and on the continental shelf. It reaches a size of nearly two metres and a weight of about 70 kg. The spawning season is uncertain but the species is known to have pelagic eggs. No data are available on commercial catches, but recently the West German expedition caught this species near the South Shetland Islands. (g) Dissostichus eleginoides. Found in Magellanic-South Georgia region at Kerguelen, Crozet and Marion islands. Reaches a size and weight rather smaller than D. mawsoni. No information about spawning or exploitation is available but may be included in the 'unspecified demersal percomorphs' category for the south-west Atlantic area. (h) Channichthys rhinoceratus. From the Kerguelen islands. Maximum length about 50 cm. No commercial catch data are available but the species is believed to have made up part of an estimated 120 000 tons taken from the Kerguelen archipelago in 1971—72. (i) Notothenia coriiceps (including A'. c. coriiceps and N.c. neglecta). Found around the Antarctic continent, Scotia Arc islands except South Georgia, Kerguelen and some other sub-Antarctic islands, and probably all other Antarctic islands except Macquarie. It spawns in May, produces large demersal eggs and grows to a maximum of 50 cm in length. No commercial catch data are available, but the recent West German expedition reported catching it at the South Sandwich Islands. (j) Chaenocephalus aceratus. Found at the Scotia Arc islands and around the Antarctic Peninsula. Reaches a length of between one-half and one metre. Information regarding spawning is lacking. No commercial catch data are available, but the recent West German expedition reported this species at South Georgia. (k) Notothenia magellanica. Found in Magellanic and New Zealand regions and at the islands lying on or near the Antarctic Convergence. It has been reported from the Ross and Scotia seas and may migrate there during the summer months. It reaches lengths of about 40 cm and spawns at the end of March at the Kerguelen islands; its eggs are pelagic. No commercial or other catch data are available. (1) Pleuragramma antarcticum. A pelagic species, circumpolar in distribution over the continental shelves. It is found as far north as the South Orkney Islands. Grows to between 20 and 30 cm. There is no information available about spawning. The fish has not been exploited. 2.6.2 Objectives Recent investigations gave an estimate of the standing stock of fish at the Kerguelen plateau of 130 000 tons (Hureau, 1974, and in press). Data on growth of some Nototheniids are available. Nevertheless, there is still a need for more information on such aspects as distribution and standing stock of the different populations and on the growth and mortality of the individuals. Consequently, more data are needed to understand the dynamics of fish populations. Therefore, the initial objectives concerning the fish component of the BIOMASS programme will be '.o concentrate on the basic studies with the following general objectives: H-34 RESEARCH PROGRAMMES Pie.udoch.a.znichthyi gzoigianm (l&ngih to so cm) Dissostiahus mawsoni (length to 2 m) Dissostiahus eleginoides (length to 1-3 m) Channichthys rhinoceratus (length, to 50 Crn) FIG 4. Antarctic fishes in the order discussed in the text. H-35 RESEARCH PROGRAMMES Notothenia aoriiceps (length to 50 Cm) Chaenoaephalus aoeratus (length to 0-5-1 m) Notothenia magellanica xjl/ (lena/th. to -W cm) Pleuragrairma antarotiaum (length to SO-50 cm) FIG 5. Antarctic fishes in the order discussed in the text. H-36 RESEARCH PROGRAMMES 1 . Stock separation and evaluation of gross potential of fish resources. 2. Development of systems for monitoring stocks that are or may become exploited. 3. To provide estimates of population parameters for models of fish population dynamics. 4. To undertake biological studies to refine the models mentioned in Objective 3 and to improve estimates of yield. 5. To develop methods for understanding, monitoring and predicting the effects of fisheries' exploitation on other elements of the ecosystem. 2.6.3 Research programmes The following specific programmes are proposed in order to achieve the above objectives: 1. Catch and effort statistics should be collected and inspected, a task which ought to be the responsibility of the government of any nation engaged in commercial fishing. The statistical areas should follow FAO standards. 2. Exchange programmes for age determination should be carried out. 3. Regular sampling for length (and, where possible, age) should be made from commercial catches. For this purpose, the length measurement used should be standardized, and the recording of total length, to the nearest centimetre below, is recommended. Efforts should also be made to collect information on the age structure (and failing this, size structure) of stocks of fish that have not yet been subjected to significant exploitation. This information should provide a unique opportunity for estimating natural mortaUty. 4. Data should be collected on the food of fishes (particularly commercial fishes) and of the principal predators, so as to quantify the predator-prey relationships which are important for trophodynamic studies. 5. Sampling of fish should be extended in areas and time as far as possible (including the use of non-commercial gear) so as to provide better information on distribution, stock separation, reproductive behaviour, possible movements and seasonal abundance. Shore stations could play an important role in this respect, and in particular morpho- logical and electrophoretic techniques should be used. In addition, such sampling will facilitate studies of reproduction and other aspects of the life history which are important for refining population models. 2.7 Cephalopods 2.7.1 Introduction Cephalopods (squids and octopuses) are known to be important organisms in the trophic structure of the Antarctic ecosystem; they constitute significant portions in the diets of Sperm Whales, seals, penguins, pelagic birds, and fishes (see Section 2.5, Tables 1 and 2). However, it is possible that they could be less abundant in the Antarctic than in the sub-Antarctic area. The highly evolved nervous system of the cephalopods allows them to be aggressive, fast-swimming predators. This characteristic, unique among invertebrates, makes them behaviourly comparable to many large predatory vertebrates. Knowledge about their biology and, indeed, about their species composition, is almost completely lacking, due primarily to the strong swimming and net-avoidance capabilities that have prohibited adequate sampling. Therefore, one of the first steps toward elucidating the biology and resource material of cephalopods must be the development of suitable fishing gear for Antarctic conditions that can catch squids in reasonable numbers even if not in a fully quantitative manner. H-37 RESEARCH PROGRAMMES Cephalopods inhabit both pelagic and benthic habitats in Southern Ocean waters. The squids are primarily pelagic forms and constitute the major resource potential in terms of numbers of species and biomass. The octopuses are primarily benthic dweUers and their biomass and consequent energy flow are relatively minor in comparison to pelagic cephalopods. This is especially the case in the Antarctic shelf waters, whereas sub-Antarctic islands appear to support larger populations of octopuses. PLATE 6. A midwatcr squid (Histioleiilhis sp); total length approximately 25 cm. Currently no commercial fishery for cephalopods exists within the confines of the Southern Ocean, but an active Japanese fishery has developed in New Zealand waters for Nototodarus sloani. Recent statistics of this fishery are as follows: Year Catch (in tons) No of boats 1972/73 13 423 71 1973/74 14 761 156 1974/75 18 947 151 1975/76 19 598 128 2.7.2 Objectives The objectives of the research programme are: 1 . To delineate the species composition of the Southern Ocean cephalopod fauna. 2. To conduct biological studies on species important to vertebrate predators and those potentially valuable as fisheries resources, both in relation to the size of the stocks and to predation rates. The proposed research will carry out basic biological studies based upon a comprehensive sampling programme which will utilize multiple types of biological and commercial collecting gear and include the stomach contents of cephalopod predators. H-38 RESEARCH PROGRAMMES 2.7.3 Research programmes 1. Biological research. (a) Extensive taxonomic research on Southern Ocean cephalopods is required before detailed biological studies can be made. A correct assessment of the population and subsequent decisions concerning the management of the stocks are dependent upon accurate identification of species and an understanding of the biology of each species. (b) Biological studies on the more important species should include distribution, vertical range, seasonal occurrence and abundance, life history, feeding strategy, reproductive potential, migrations and relationships to environmental factors. (c) In order to utilize fully the limited cephalopod material available, research programmes need to take advantage of every potential source of material, such as from krill trawling operations, or predators' stomach contents. (d) Studies should be conducted to determine the nutritional qualities of different species and even of different parts of the animals (eg, mantle v viscera) to assess their value to predators and to human consumers. (e) Research needs to be conducted, perhaps at laboratories outside the Southern Ocean area, on the behavioural responses of cephalopods to determine the effectiveness of various catching devices. (f) Attention should be given to rates of predation as well as to the size (biomass) of the cephalopod stocks. 2. Sampling techniques. The paucity of knowledge about cephalopods is directly related to problems of adequate sampling. Therefore, in order to conduct basic biological studies, we propose that: (a) A wide variety of trawling techniques be utilized, including traditional mid-water trawls (eg, 3m IKMT and RMT 1&8) and large commercial trawls (eg, Engel's trawl), and that trawUng with these large nets be conducted at depths greater than is customary for krill operations, ie throughout the water column and especially at the hitherto unsampled bentho-pelagic zone near the bottom. (b) Stomach contents from known predators (particularly Sperm Whales, seals, penguins, pelagic birds and fishes) be utilized for sampling cephalopods. A study of cephalopod beaks for identification and biomass assessment should be initiated based on material from identified cephalopod specimens. Eventually this should lead to the abiUty to identify beaks alone, which are so frequently found in predators' stomachs. A study of beaks and statoliths taken by deep benthic dredges should aid in assessing distributions of species. (c) Larval collections be conducted utilizing plankton, neuston and micronekton nets and used as an indirect means to assess cephalopod populations. (d) Acoustic techniques be developed for locating aggregations of cephalopods and used in association with trawling operations. (e) Techniques other than pelagic and benthic trawls be employed to explore alternative methods for sampling and assessing cephalopod populations, for example, large purse seines, night lighting and jigging, baited cameras, submersibles with side-scanning sonar, TV and camera-equipped sleds. (f) A standardized sampling protocol be established to allow comparison of results from various programmes, such as depth regime, or catch per unit effort. H-39 RESEARCH PROGRAMMES (g) Net closing devices be used to ensure precision in determining vertical distributions of species. 2.7.4 Implementation of research programmes If the objectives of the proposed research programme on cephalopods are to be achieved, full advantage must be taken of the several programmes of implementation. Because of the unique situation of cephalopods in the Antarctic ecosystem, it is proposed that the delineation of the fauna and an understanding of its biology can be most effectively achieved by conducting a separate research project. Several ships will be equipped with standardized sampling and acoustical gear to ensure direct comparability of results. The proposed cephalopod research programmes can be conducted in collaboration and co-operation with other biological projects involved in the multi-ship programme. In addition, much valuable data can be obtained from participation in both the supporting ship-based programme and the supporting shore-based programme. In particular, co-operation with commercial trawlers and whaling factory vessels should provide specimens and data that otherwise could not be efficiently obtained. SELECTED REFERENCES ANDRIYASHEV, A. P. 1965. A general review of the Antarctic fish fauna. In: OYE, P. van, and MIEGHAM, J. van, eds. Biogeography and ecology in Antarctica. The Hague, W. Junk, p 343—402. Dewitt, H. H. 1971. Coastal and deep water benthic fishes of the Antarctic. Antarctic Map Folio Series, Folio 15. EvERSON, I. 1970. The population dynamics and energy budget of Notothenia neglecta Nybelin at Signy Island, South Orkney Islands. British Antarctic Survey. Bulletin, No 23, p 25-50. HuREAU, J.-C. 1970. Biologie compaiee de quelques poissons antarctiques (Nototheniidae). Bulletin de I'Institut Oceanographique (Monaco), Vol 68, No 1391. HuREAU, J.-C. 1974. Les possibilites d'exploitation des resources marines dans lestles australes fran^aises. Bulletin du Museum Natioruil d'Histoire Naturelle (Paris), 3^ serie. No 154, p 185-91. HuREAU, J.-C., in press. Caracteristiques generales de la faune ichtyologique du secteur indien de I'ocean antarctique et estimation du stock de poissons autour des lies Kerguelen. Memoires du Museum National d'Histoire Naturelle (Paris). 2.8 Benthic invertebrates of potential commercial importance 2.8.1 Introduction Although research on the composition, metabolism and productivity of the benthos is needed to improve our understanding of the structure and processes of the Antarctic marine ecosystem (Mills, 1975), in this section the research programmes will be directed primarily towards those organisms which are of potential commercial importance. This, as was stated earlier (Section 2.1), does not imply that research on benthic communities is unimportant, merely that detailed study of these communities is beyond the immediate scope of this report. For benthic invertebrate resources, the following islands were considered in addition to the shelf of the Antarctic continent: H-4 0 RESEARCH PROGRAMMES Latitude 30°-40°S 40°-50°S 50°-60°S 60°-70°S (a) Rock lobsters (spiny lobsters). Islands St Paul and Amsterdam Tristan da Cunha Marion and Prince Edward Croze t Kerguelen Antipodes Bounty Gough Bouvet Heard Macquarie Auckland Campbell South Georgia South Sandwich All the islands south of 60°S Plate 7. Rock lobsters (Jasus sp). In the Southern Ocean no benthic invertebrates are commercially harvested. However, within the wider area considered here, rock lobsters of the genus /axux are currently exploited to a marked extent. At the Tristan group of islands (Tristan, Nightingale, Inaccessible, and Gough) and at Amsterdam/St Paul they have been exploited for a number of years, and for both groups of islands catch and effort statistics are available. The size composition of the catch and the production by tail size category have been recorded. Fairly good data on growth of rock lobsters at the Tristan group have been obtained from tagging experiments, which are to be continued. Analysis of size composition data has been attempted to evaluate growth rates at Amsterdam/St Paul, but the results need verification by other techniques. H-41 RESEARCH PROGRAMMES Rates of tagging at Tristan and neighbouring islands have been insufficient to provide good estimates of mortahty. Stock sizes and mortahty have been calculated from catch per unit effort information at Amsterdam and St Paul. Estimates of stock range from 7 000 to 14 000 tons (Vranckx, 1974). Catch per unit effort trends, as well as catch size composition, suggest that stocks are not over-exploited at Tristan and adjacent islands, and that present catch levels of about 800 metric tons nominal weight can be maintained without increasing fishing effort (Pollock, 1976). Similar analysis at Amsterdam/St Paul has indicated that stocks have decUned, and the yield of 900 metric tons obtained in the past may be too high (Vranckx and Hureau, in press). (b) lithodid crabs. In recent years, exploratory fishing surveys have been conducted by France to collect biological information on and to assess the potential of lithodid crabs, particularly Lithodes murrayL Experimental fishing resulted in large catches around the Crozet islands. Few crabs were caught at Prince Edward Island and catches were negative at Marion and Kerguelen islands (Amaud and Do Chi, 1976). It was felt that further surveys were required to evaluate completely the potential of these islands. In the course of the surveys, biological observations on reproduction, parasites and feeding were also made. PLATE 8. A lithodid crab (Lithodes murrayi). (c) Spider crabs. The spider crab Jacquinotia edwardsii is endemic to southern New Zealand and the New Zealand sub-Antarctic (40°-50°S, 100°-165°E). It has been recorded in decreasing quantities at Pukaki Rise, Auckland Islands, Campbell Island, Bounty Islands (Anon, 1971), Stewart Island Shelf, Puysegur Bank and off the east coast of the southern South Island. The presence of large stocks in depths of less than 200 m at the Auckland Islands has also been shown (Ritchie, 1970, 1973; Ryff and Voller, 1976). Experimental fishing by a joint New Zealand-Japanese expedition during the austral summer found a resource at the Pukaki Rise which was about H-42 RESEARCH PROGRAMMES three times that found at Auckland Islands: estimates of possible annual harvest ranged from 385 000 to 530 000 crabs (Ryff and Voller, 1976). Apart from the stock estimates mentioned above, there is information on experimental fishing and processing, and some brief observations on moulting, reproduction, feeding and behaviour. Generally there is a paucity of information on the biology and population dynamics of this species. (d) Other benthic invertebrates of potential commercial interest. Other crustacean species, which have little potential value in view of their low population numbers and/or difficulty in catching are: the Antarctic shrimps {Notocrangon antarcticus and Oiorismus antarcticus), other species of lithodid crabs especially Paralomis sp, and the smooth red swimming crab Nectocarcinus bennetti. Except for the scdWo^p Adamussium colbecki and tlie clam Laternula elliptica, the Antarctic shelf appears to contain few species of bivalve fauna similar to those exploited elsewhere. The sub-Antarctic islands also lack such bivalves, except for Mytilus ediilis desolationis and Aulacomya ater which are abundant at Kerguelen, and M.e. desolationis and A. maoriana around the New Zealand sub-Antarctic islands. Several other benthic invertebrate species are possibly suitable for human consumption. These include the gastropod Neobuccinum eatoni and sea urchins of the genus Sterechinus. They occur in shallow water and are generally circumpolar in distribution. In view of the small size of the stocks and the logistic problems involved, it w concluded that the harvesting of these resources is not practical. 2.8.2 Objectives The research programme for benthic invertebrates is to be established within the framework of the following broad objectives: 1 . Assessment of the state of presently exploited stocks. 2. Identification of new resources and assessment of their magnitude. 3. Establishment of monitoring systems for exploited stocks. 4. Evaluation of biological and environmental factors which will lead to refinement of population models. 2.8.3 Research programmes Assessment of the potential of unexploited stocks requires systematic fishing, for which the following proposals can be made. 1. Catch and effort. The distribution of both rock lobsters and crabs is usually closely related to depth. PreUminary investigation of bathymetry, or acoustic surveys, can facilitate the systematic setting of traps and the delineation of areas of interest. A preliminary gross estmiate of stock size can be made by combining catch rate with survey area and depth information. Catch and effort statistics should be collected; these should include data on numbers, nominal weight, and production weight, as well as number of vessels, type of gear, number of traps per vessel, and number of trap days, hours or hauls. The nature of the catch and/or production should be recorded by data on size composition, state of exoskeleton, sex ratio, state of reproductive organs, and size at maturity. It is recommended that rock lobster size be expressed as carapace length, measured from the tip of the rostrum to the mid-dorsal end of the carapace, and that the size of lithodid crabs be measured from the posterior orbit to the mid-dorsal end of the carapace. H-43 RESEARCH PROGRAMMES 2. Growth. This is difficult to measure in decapods. Tagging and recovery after mouUing and short term holding experiments can provide good data on growth of adults. Polymodal frequency analysis can indicate growth rates of juvenile specimens. 3. Mortality. Size composition data combined with growth information may provide rough estimates of total mortality, or natural mortality if size composition information is collected in the early stages of fishing. The rate of decline in catch per unit effort, during a short fishing season, in relation to the catch, can also provide an estimate of total mortality. The above techniques require that catchabiUty remains constant with size and throughout the fishing season. Tagging experiments probably provide the most promising means of estimating fishing mortality. Tag loss is usually confined to the moult. 4. Yield assessments. Trends in catch size composition and catch per unit effort reflect the impact of fishing on the stock. Better yield assessments can be made from stock production curves if a suitable time series of catch and effort data is available. If growth and mortaUty data are available, yield per recruit curves can be calculated to evaluate the sensitivity of the stock to changes in effort and sizes at first capture. The above proposals refer mainly to stock assessment needs. Biological studies which would enhance population models and management should encompass the following: seasonal migrations; stock separation; feeding and predation; and larval dispersion, and recruitment. SELECTED REFERENCES Anon. 1971. Kaiyo-Maru's survey report - New Zealand southern waters - for the year 1910. Japanese Fishing Agency, unpublished report. Arnaud, p. and Do Cm, C. 1976. Proposals for the study of living resources of the Southern Ocean: the Lithodidae (Crustacea, Anomura). UnpubUshed report. Mills, E. L. 1975. Benthic organisms and the structure of marine organisms. Journal of the Fisheries Research Board of Canada, Vol 32, p 1657-1663. Pollock, D. E. 1976. Assessment of the rock lobster (Jasus tristini) stocks. Cape Town, Sea Fisheries Branch, unpubUshed report. Ritchie, L. D. 1970. Southern spider crab {Jacquinotia edwardsii) at the Auckland Islands, October 1971. Fisheries Technical Report, No 101. Ritchie, L. D. 1973. Commercial fishing for southern spider crab (Jacquinotia edwardsii) at the Auckland Islands, October 1971. Wellington, Ministry of Agriculture and Fisheries. (Fisheries Technical Report No 101.) Ryff, M. R and VoLLER, R. W. 1976. Aspects of the southern spider crab (Jacquinotia edwardsii) fishery of southern New Zealand islands and Pukaki Rise. Fisheries Technical Report , No 143. VrANCKX, R. 1974. Evolution du stock de langoustes sur les fonds de peche des Ties St Paul et Nouvelle Amsterdam de 1962 a 1970. Bulletin du Museum National d'Histoire Naturelle (Paris), No 155, p 193-204. Vranckx, R and HuREAU, J.-C, in press. Production de langoustes dans les eaux des Ties St Paul et Amsterdam. In: DUNBAR, M. ed. Polar oceans. Calgary, Arctic Institute of North America. 2.9 Seaweeds 2.9.1 Introduction The littoral zones of the Antarctic and sub-Antarctic are particularly rich in benthic seaweeds. About 700 species belonging to 300 genera have been recorded from the Southern H-44 RESEARCH PROGRAMMES Ocean (Delepine, 1966 and in press; Neushul, 1968; Zaneveld, 1966). There are representatives of 56 genera that have been utilized either as food or for the production of algal products in many parts of the world. Worldwide the utilization of seaweeds is constantly increasing. Therefore, it can be expected that attention in the future will be paid to the seaweed resources of the Antarctic and sub- Antarctic. UtiUzation may take the form of human food, fodder for domestic animals, or the production of agricultural fertilizers, phycocoUoids such as alginates, agar-agar and carrageens and medicinal products such as antibiotics. It should be stressed that benthic marine algae are an important component of the coastal ecosystems where they contribute significantly to the overall primary production. They also provide a substantial food-source for many fishes and invertebrates, either directly or through the detritus pathway, as well as a substrate and shelter for a wide range of animal species. In addition, algae are known to excrete various substances in large quantities, but the role of these substances in the ecosystem is not yet fully determined. Species of immediate interest in the Southern Ocean are the brown algae Macrocystis pyrifera and Durvillea antarctica and the red algae 'Agarophytes' such as the Gelidiales or Gigartinales (Gracilaria. Gigartina sp, etc). Some of the red algae are currently being exploited in South America, South Africa and New Zealand. Quantitative information has been obtained on Macrocystis pyrifera from the South Atlantic islands, and especially from Kerguelen, where it is estimated that the beds cover more than 200 km^ , with biomasses ranging from 3.4-22.5 kg/m^ with means of 5-10 kg/m^. On New Zealand shores the standing stock of Durvillea antarctica ranges from 10-20 kg/m^ . Standing stocks of a similar order of magnitude have been recorded from the Kerguelen islands. 2.9.2 Objectives Research objectives are: 1. To gain an understanding of the role of benthic algae in coastal ecosystems of the Southern Ocean. 2. To carry out basic studies on the potentially exploitable benthic algal species in the Southern Ocean, and to assess the importance of this resource. 2.9.3 Research programmes 1. Stock estimation. (a) Mapping of the distribution of selected species. The use of satellite imagery for this purpose should be explored. (b) Estimation of standing stock and primary productivity in selected habitats. Such estimations should be carried out at different seasons and combined with estimates of total standing crops for the species studied. (c) Resettlement and growth rates after clearing or harvesting in order to obtain information on the subsequent history of the cleared or harvested areas as related to the ecophysiology of the colonizing species. (d) Study of the vertical distribution of the algal standing crops. 2. Ecophysiological studies. (a) In situ (phenological studies): to include studies on growth rates, productivity, longevity, intensity of reproduction related to environmental factors such as light, photoperiodicity, temperature, etc. (b) In the laboratory: studies to determine the ecophysiological reactions to environmental factors for each phase of the life cycle; life cvcle studies. H-45 RESEARCH PROGRAMMES 3. Biochemical studies. Biochemical composition with season and age, including estimation for each period of the year of the quantities of different chemical constituents (nitrogen, phyllocolloids and other substances such as antibiotics). 4. General ecological studies. (a) Studies of the role of algal species in the ecosystems, their utilization by consumers and their relationship to fishes and invertebrates. (b) Life cycle studies both in the field and the laboratory. (c) Taxonomic studies based on data from populations in the field. (d) Long-term standardized measurements of environmental factors such as radiation, temperature and ice cover. SELECTED REFERENCES DelEPINE, R. 1965. La vegetation marine dans I'Antarctique de I'Ouest comparee a celle des lies Australes l'"ran(;aises: consequences biogeographiq ues. Comptes rendus des seances de la Societe de biogeographie , Vol 374, p 52-68. DelEPINE, R., in press. Seaweeds of the Antarctic and sub-Antarctic. In: EL-SAYED, S. Z. ed. BIOMASS Vol 2: Selected contributions to the Woods Hole Conference on Living Resources of the Southern Ocean, 1976. Cambridge, Scientific Committee on Antarctic Research. NeuSHUL, M. 1968. Benthic marine algae. /InfarcncAfop foto Series, Folio 10, p 9-10. ZanevelD, J. S. 1966. Vertical zonation of Antarctic and sub-Antarctic benthic marine algae. Antarctic Joumalofthe United States, Vol 1, p 211-13. 2.10 Remote Sensing 2.10.1 Introduction Remote sensing is a technique of growing importance in marine science. The remote sensing systems can take the form of devices used from aircraft, or satellite platforms, or ocean data buoys. Aircraft-borne sensors can possess different characteristics from sateUite sensors. Thus, each hardware system can provide, to a large degree, different types of information; and studies and/or experiments must be designed with these different capabilities in mind. 2.10.2 Objectives Remote sensing techniques can potentially be used to assess the abundance of phytoplankton, macro-algae, krill, birds, and mammals. The application of remote sensing is briefly reviewed in the following sections. 1. Phytoplankton. In general there is a positive correlation between the phytoplankton standing crop (in terms of chlorophyll a) and primary production of surface waters; hence measurement of chlorophyll levels may be useful in assessing the productivity of these waters (El-Sayed, 1971). In recent years, there have been some pioneering studies of remote sensing of chlorophyll a, temperature, etc, using satelhtes and low flying aircraft (El-Sayed and Green, 1974). An excellent opportunity to contribute to our knowledge of using remote sensing techniques to study the standing crop of phytoplankton and other biological and physical attributes of the water column will soon be available through the use of the Coastal Zone Colour Scanner (CZCS) on the NASA NIMBUS-G sateUite to be launched in 1978. It is interesting to point out H-46 RESEARCH PROGRAMMES in this respect that one member of the Group of Specialists (S. Z. El-Sayed) is also a member of the CZCS NIMBUS-G Experiment Team. 2. Krill. The activity of euphausiid swarms produces a long-lasting luminescence, and will probably be detectable by Low Light Level Television. The images from such sensors may be stored on videotape for later processing. Thus, the night-time aerial/acoustic method of abundance estimation which has been developed for pilchard (Cram and Hampton, 1976) may be appUcable to that part of the krill population occurring near the surface in ice-free areas where sufficient darkness can be expected. Alternatively, if the visibihty of shoals is adequate, the numerous daylight photographic and spectro-radiometric techniques reviewed by Benigno (1970) miglit also be highly suitable. Krill swarms have been recorded to have colour characteristics ranging from brilliant red to ochre and yellow. Such variation in colour probably indicates differing biological character- istics; hence, for successful identification with remote sensing, baseline work is needed to interpret the relationship between colour and biology of the species. 3. Birds and mammals. This group is most visible on the surface of ice or on land but less conspicuous when swimming in the water. Each group and/or species have characteristics which make them unique when considering remote sensing applications. The penguins congregate on rookeries to breed and nest. During this time their density is high, and concentrations of over 100 000 animals are found. However, the seals of the region rarely concentrate in large numbers and are usually distributed in small groups. Exceptions are the WeddeU, Elephant and Fur seals, which congregate yearly to breed. It appears that the cunent instrumentation of satelHtes is of Umited use for the direct sensing of vertebrates of the region because its resolution is limited. The vertebrates individually or in concentrations are probably of insufficient size and show insufficient contrast in emissivity to be detailed by present satellite instrumentation. Thus, aircraft sensors will probably be needed. The current aircraft which are available are limited in range; consequently, such surveys may present a difficult task. 4. Land, ice, or sea-based sensors. Another type of remote sensing that may be particularly important is the use of remote sensors placed at the surface, which allows information to be transferred to ground base or sateUite base data systems. This capability would be extremely valuable in the Southern Ocean, where access during the winter months is nearly impossible. Static platforms could be located in coastal regions with links to sensing systems in the water; alternatively, drifting sensing systems could be set adrift in the pack ice regions to monitor physical and biological parameters, as well as to pick up acoustical indications of seals, whales, and penguins. In summary, the need for remote sensing in the Antarctic regions varies considerably depending on the species involved and the particular problem under study. However, it is clear that if we are going to evaluate and monitor successfully the biological resources of the Southern Ocean, it is essential; (a) to look to these techniques in the future, and (b) to begin to develop them now so that they may be adapted to the problems and situations unique to species of the Southern Ocean. 2.10.3 Research programmes I. Existing and near future sateUite data and instrumentation. LANDSAT 1 has obtained three and one-half years of data on ocean turbidity which could be utilized in any pre-operational study. LANDSAT 2 was launched in 1975 and is working with much the same capabilities as LANDSAT 1. Unfortunately, the LANDSAT 1 imagery of H-47 RESEARCH PROGRAMMES the Antarctic is limited to south of approximately 65°S except for certain runs up the South American coast, which may render these data of little value. Nevertheless, the resolution is 75 X 56 m and sequential coverage is available every nine days. LANDSAT 2 has a similar coverage but could acquire data from other areas on command. If data are required from the sub-Antarctic islands, for example, a request to NASA for this coverage would probably succeed if the motivation were adequate. Information on the extent of likely cloud contamination of satellite data can be obtained from the Temperature Humidity Infra-Red Radiometer Systems on NIMBUS satellites, and from the visible and infra-red systems on NOAA satellites. The extent of ice cover can be determined from the scanning microwave radiometer on N0AA5 and, after launch, from NIMBUS-G. The US Naval Weather Office produces maps of ice distribution in three-day averages. It is possible that such maps may be made available to scientific organizations. 2. Feasibility study of krill by remote sensing. As a first step in the development of a krill remote sensing programme, a feasibility study should concentrate on acquiring imagery of krill swarms with currently operational sensors and at low cost. Therefore, the spectral and size characteristics of krill swarms should be measured as primary data for sensor specifications. Photographic spectro-radiometers and Low Light Level Television equipment should all be used. The feasibility study should concentrate on the Antarctic Peninsula area in middle to late summer, and will require a long range small to medium sized aircraft with a fuel capacity of 10-12 hours flying time. High precision navigation equipment is essential, together with radar, a radar altimeter and standard camera hatch for sensor mounting. It is estimated that 50 hours mission rime (total 100 hours) would be adequate for the feasibility study. Operations would be most convenient from bases on the southern tip of South America or in Antarctica. 3. Feasibility study of mammals and birds. (a) Density and distribution studies by passive sensors. At present no specific programme can be outlined, but it is important to initiate trials of various sensors such as water penetration film, running television systems and laser systems. (b)Sensors for biological measurements of instrumented animals. The development of instrumentation and monitormg stations to measure body temperature, heart rate and other physiological characteristics should be initiated. Techniques are well established for terrestrial vertebrates and circuitry is well advanced. The feasibility of using ocean buoys to relay data from instrumented animals to satellites should be considered. With larger animals it might be possible to transmit directly to sateUites. SELECTED REFERENCES BeNIGNO, J. A. 1970. F'ish detection through aerial surveillance. Madern Fishing Gear of the World, Vol 3, p 44-48. (Fishing News (Books) Ltd, London.) Cram, D. L.and HaMHTON, I. 1976. A proposed aerial/acoustic strategy for pelagic fish stock assessment. Journal du Conseil permanent international pour I't'xploration de la Mer, Vol 37, No 1. El-SaYED, S.Z. 1971. Dynamics of trophic relations in the Southern Ocean. /n: QUAM, L.ed. Research in the Antarctic. Washington, DC, American Association for the Advancement of Science, p 73-91 . El-SayED, S. Z and Green, K. A. 1974. Use of remote sensing in the study of Antarctic marine resources. In: BOCK, P.. BAKER, F. W. G., and RUTTl NBl RG. S. eds. COSPAR approaches to earth survey problems through use of space techniques: proceedings of the symposium held in Constance . . . 1973. Berlin, Akademie-Verlag, p 47-63. I VANOV, B. G. 1969. O svechenii antarkticheskogo krilya (b'upliausia superba) |Lumincscencc of Antarctic krill (t'uphausia superba)] . Okeanologiya . Tom 9, Vypusk 3. p 505-06. H-48 RESEARCH PROGRAMMES Mark, J. W. S. 1962. The natural history and geography of the Antarctic krill (Eupliausia superba Dana). Discovery Reports, Vol 32, p 36-434. MOISEYEV, P. A. 1 970. Some aspects of the commercial use of the krill resources of the Antarctic seas. In: HOLDGATE, M. W ed. Antarctic ecology. Vol 1. London and New York, Academic Press, p 213-16. SiNIFF, D. B. and KeucHLE, V. B. 1974. Remote sensing of Antarctic biological resources: vertebrates. In: BOCK, P., BAKER, F. W. G., and RUTTENBERG, S. eds. COSPAR approaches to earth survey problems through use of space techniques: proceedings of the symposium held in Constance . . . 1973. Berlin, Akademie-Verlag, p 65-74. Staples, R. F. 1966. The distribution and characteristics of surface bioluminescence in the oceans. Washington, DC, USN Oceanographical Office. (Technical Report TR-184.) H-4 9 3. PRACTICAL IMPLEMENTATION OF THE RESEARCH PROGRAMME The objectives of BIOMASS require action along the following Unes: 1. Seagoing operations of research vessels. These should include both detailed multidisciplinary studies in limited areas of special interest (co-operative 'experi- ments'), and large scale surveys which will build on the work carried out by such research vessels as the Discovery, Ob and Eltanin and by recent exploratory cruises. 2. Other operations at sea. Exploratory and commercial fishing and whaling vessels, supply vessels and others are expected to be operating in the Antarctic during the BIOMASS period. Observations from these vessels will be used to supplement the work of the research vessels, especially in relation to the harvestable resources. Seagoing operations should be supported by remote sensing work. 3. Shore-based studies. Investigations on feeding, growth and reproduction of krill and other key organisms at established coastal stations should complement and add year-round continuity to the offshore ship-based studies. 4. Data analysis. Arrangements will be made to ensure the proper compilation, storage, dissemination and analysis of information arising from all relevant activities in the Antarctic. 5. Modelling. Immediate use will be made of information from published and unpublished sources for the development of models; field and experimental data from (1), (2) and (I), (2) and (3) will be used to expand the models as they come in. 3.1 Seagoing activities of research vessels During the first operational stage, culminating in the First International BIOMASS Experiment (FIBEX), 1980-81, investigations should be concentrated in a suitable area such as the Scotia Sea or the Atlantic sector in general. It is in this sector that the resources appear most abundant and where they are most likely to be harvested. Studies of other sectors must follow. 3.1.1 Macroscale studies of krill in relation to hydrography The drift of krill during its two to three years of life is of the order of 1 000 km, and is mainly governed by surface and deep currents. The assumed 'developmental ascent' may include the upper 2 000 m of the water column and, together with larval development, takes between 10 and 100 days. Two proposed projects are sufficiently wide ranging to provide information on transport mechanisms: the Weddell Sea Gyre project, and the International Southern Ocean Studies (ISOS) project on water circulation and remote sensing of the variability in oceanic fronts and ice movements. The near surface distribution of krill in summer has been described in considerable detail by earlier expeditions. However, little is known of the transport of the various life history stages and of the Unks between the different populations. Quantitative sampling of krill at several depths should be carried out in various areas and seasons: there is an urgent need to extend the life history studies of krill into autumn and winter and into the pack ice zone. The sampling programme should be guided by oceanographic observations and supported by radio echo surveys. At least two research vessels will be needed for each of the selected areas. H-50 PRACTICAL IMPLEMENTATION OF THE RESEARCH PROGRAMME 3.1 .2 Microscale studies on the ecology ofkrill swarms The interaction between phytoplankton composition and biomass and the swarming of krill under the influence of oceanographic factors can be studied at the rate of 10 km per 10 days; two to three ships are needed in a complete experiment of this type. Measurements to provide information on vertical transport and stability and on advection should be made, and samples taken in order to determine the spatial distribution and densities of swarms of krill in relation to the abundance and size spectra of phytoplankton, which will enable us to understand the reactions of a krill swarm to its food base and the effects of grazing on the phytoplankton. Quantitative recordings and sampling for analyses of phytoplankton distribution and zooplankton abundance and composition in different depth layers and at different times of day and night will provide additional information on the spatial structure of the biotic environment of krill. Echo sounding and sonar observations should provide three-dimensional pictures of the changes in time of zooplankton concentration. These studies should be carried out at several selected places, mainly in areas of high kriU concentration at the ice edge, in the neritic zone of the Antarctic Peninsula and of the Antarctic islands, as well as in the open ocean of the West Wind Drift, the East Wind Drift and the upwelling zones. If possible the studies should be repeated at different times of the year. 3. 1 .3 Microscale studies of food chain structures and functions In order to learn more about possible future effects ofkrill fishing on the structure of the first and second trophic levels, the composition of phytoplankton and its productivity should be studied in places and seasons which differ in relative abundance of krill, salps or other herbivores. These studies must be based on quantitative plankton sampling at various depths, and can be considered as an extension of 2.2.2 into areas and seasons where alternative grazing and predation strategies lead to changes in the quantitative composition of the herbivore level. 3.1 .4 Macroscale studies of zooplankton, nekton and benthos There is still a need for further quantitative sampling of zooplankton, pelagic squids and fishes in various regions of the Southern Ocean, and the benthos of the Antarctic is poorly known. Quantitative data are needed on the abundance of the dominant elements of plankton and benthos in the different regions. The position of the dominant species within the food chains should be ascertained by studies of stomach contents and feeding apparatus. Benthos studies may be combined wath observations on sedimentation and decomposition of fecal pellets, dead zooplankton, particularly krill, and diatoms. The biological and sedimentological surveys are very time consuming, and more than one research vessel will be needed, even if the intention is to cover only one oceanic sector each summer. 3.1.5 Seagoing activities of other vessels The seagoing research tasks require the co-operation of a number of oceanographic and other vessels in the Southern Ocean. The majority of the seagoing studies need fully equipped research vessels, particularly the multidisciplinary studies on the relationship between krill and the environment. The success of the biological programmes will depend largely on good design and effective integration with physical studies of horizontal transport and vertical mixing. It would neither be sufficient nor economical to add small biological programmes to physical expeditions and vice versa, but full scale programmes of both disciplines could be combined. International co-operation is essential in view of the large support needed for a multidisciplinary study which includes several vessels and a considerable number of speciahsts. H-51 PRACTICAL IMPLEMENTATION OF THE RESEARCH PROGRAMME 3.2 Other ship-based programmes The work of the research vessels can be greatly assisted by the co-ordinated survey efforts of exploratory and commercial traviflers, the few whaling vessels which are still in existence, and by supply ships and icebreakers. A brief summary of the advantages and limitations of the different types of vessels with regard to tlie research objectives of BIOMASS is given in Table 4. Given the ambitious objectives of BIOMASS, its success will depend on making the best use of all types of vessel. In view of the importance given to resource management, there will be special emphasis on obtaining information from those vessels most directly concerned with the harvestable stocks, ie, exploratory and commercial vessels, particularly those directed at krill. This information will include detaUs of the catch, the distribution and, as far as possible, the relative abundance of the stock, all of which are important in studying the population dynamics of the exploited stock. In addition, where opportunity offers, other biological observations using simple equipment such as plankton nets will be made from these and other non-research vessels. Information from fishing vessels is, however, no substitute for observations from research vessels. Indeed, to the extent that the number of the former increases— in other words, exploitation intensifies-there will be a greater urgency for timely scientific advice on management, some of the information for which is only obtainable wath well equipped research vessels. 3.2.1 Utilization of ships 1. Exploratory trawlers. At present, exploratory trawling is being carried out in the Southern Ocean by ships from the Soviet Union, Japan, the Federal Republic of Germany and Poland, and this activity is likely to increase in the future. Each exploratory trawler has, at present, an active research programme, but these are being conducted largely in isolation from one another. This fleet of trawlers constitutes the largest single potential for supporting the proposed BIOMASS programme. Although the spectrum of opportunities available for oceanographic studies on such vessels is likely to be less than that on research vessels, their unique capacity for effective sampling of krill en masse and their sophisticated hydro-acoustic technology should be fully utilized. Successful liaison with the agencies operating these vessels would not only enhance the significance of their observations relative to one another, but would also provide 'ground truth' for other operations, such as acoustic surveys by other classes of vessels operating in the Southern Ocean, and remote sensing of surface swarms of krill. Exploratory trawlers (and commercial trawlers) have great potential for making significant contributions to our knowledge about krill swarms by taking simple qualitative samples of phytoplankton (mesh size 35 /im) and zooplankton (mesh size 200 Aim). 2. Commercial trawlers. At present, most of the commercial trawlers in the Southern Ocean operate in tandem with exploratory trawlers or research vessels, but their operations in the future could well be independent. Because these trawlers are dictated by considerations other than research, the spectrum of opportunity which they provide for BIOMASS is more limited than that provided by exploratory trawlers. However, the success (or failure) of their trawling operations and the gathered acoustic observations constitute data of prime importance if they can be extracted and integrated with those of other commercial trawlers. It would also be possible (with httle extra effort) for the facilities of such trawlers to be extended in directions they might not normally cover, for example, acoustic surveys for squids as well as krill. The BIOMASS programme should also include plankton sampling at every trawling station by means of the simple free fall net. H-52 PRACTICAL IMPLEMENTATION OF THE RESEARCH PROGRAMME CO H < o . 4) _r OT o u « n 2 ' o "^ s ;; " « p o — as oii-gi a:£ u 11 n S g a. o a: (K CO H Z as H CO Z o u Q Z < a D H ca Z < P z D H a! g 0. O S o a: < w in U OS b4 O >" < s s •t w a < Q i <<« eio HZ 5° >• Oa: £* o< w CO Z O H < H CO U o: o X CO CO _] (I) CO CO u > I u < u CO U] as o a S' £■1 25 C 3 O Urn 2 > 0. u t =1 i 2 M 4* ^ - c " i> ^ 2a 5 " 2 "t-o O O 3 = - M M n lis o S u £ o. 5! =^ o t "> w — '~ 5 " .2 O ra ** r- M (U a> tfl ^ O c o y a c 4> = :: s ° c .2 o'3 5 g c t U 4> 5 <-» c O O M o ■a .3 XI 4> D. o. n c >2 o «, £ tn ^ 3 z < as H CO Z O o 3 S u. •5R i! 2 u ■- •- u .ti -Joe ->, o JO o n e5! Z o c 3 O = w • c u -1 «i ^ -1 ** _ - .£> = 2 = ■-on: a. t« 2? £ - C.5 >. 3 I It E h-| li i: o £c 2 Is P i E a> o jf a « n w u ■o ao i> c 1 * X T3 3 c CO c B _3 1) c c 3 a B i> 'o 0 U 3 M n .2 (J u ^ w s Qfi ■a a OJ 3 (« ■5 -U 0 i> a> 0 c ■o 0 D. ^ £ « s < ■c OQ c H-53 PRACTICAL IMPLEMENTATION OF THE RESEARCH PROGRAMME 3. Whaling ships. There are still whaling vessels catching baleen and Sperm whales in the Antarctic. In addition to the information on whales that can be obtained from the activities of these vessels, they provide excellent opportunities for studying the animals eaten by the whales. No present fishing or research gear is as effective in sampUng squid as the Sperm Whale. High priority should be given to making arrangements with the whaling companies for squid biologists or technicians to accompany their ships and to collect squid material. Biological observations should also be made of other animals (krill, fish, etc) eaten by whales. It would be particularly useful to determine whether the krill eaten by whales are similar in, for example, size or age to those caught by trawlers. 4. Icebreakers. Much of the biological activity in the Southern Ocean occurs under the pack ice and to reach a proper understanding of the ecosystem one must take this into account. There is thus an urgent need for biological and also physical and chemical sampling beneath the ice. This work could best be accomplished by an icebreaker, fully equipped for oceanographic work and capable of biological sampling in deep water. 5. Supply ships. Although constrained by time, the supply ships have great potential for adding to our understanding of the position of the polar front due to their widely spaced and fairly regular tracks. They can further assist by collecting data on temperature, salinity and chlorophyll. In this way, the supply ships can provide a broad network of ground truth for remote sensing by the NIMBUS-G Coastal Zone Colour Scanner, which will be providing simultaneous coverage over a broader area (See Section 2.10). It is recommended that the national Antarctic agencies operating supply ships be invited to take the major responsibihty for this part of the BIOMASS programme. It would be useful if a working group were formed of those scientists from each national Antarctic agency responsible for the programme, representatives from the NIMBUS— G remote sensing team, and individuals experienced in continuous monitoring and processing of surface data. The second role which supply ships, and icebreakers, could play in the BIOMASS programme is the deployment of drifting sensors in satellite communication with shore stations across the Southern Ocean. The technology exists for buoy and satellite communication (Creswell, 1976), for biomass estimates in the water column (Beamish, 1971), and for measurements of turbidity or transparency. A potentially powerful means could therefore be developed within the one buoy assembly for continuous and relatively inexpensive monitoring of krill swarms. It is recommended that the development of this technology be supported by BIOMASS and implemented as soon as possible. The supply ships, and particulariy the icebreakers, can further contribute to the BIOMASS programme by making observations on plankton over a period of days and possibly weeks around and in the pack ice. The Japanese supply ship Fuji is currently carrying out such a programme off Syowa station. These observations should be integrated with tliose of a similar nature carried out on platforms of floating ice from the shore stations. 3.2.2 Recommendations 1. That national agencies responsible for the operation of support cruises for scientific bases on the Antarctic continent be invited to execute a programme of underway surface observations of temperature, salinity, and chlorophyll in the Southern Ocean, together with the collection of XBT and underway echo traces from the upper 100 m; and that a representative from the NlMBUS-G CZCS survey be invited to join tlie working party to provide liaison with the remote sensing survey of surface properties. H-54 PRACTICAL IMPLEMENTATION OF THE RESEARCH PROGRAMME 2. That a working party be established to plan and execute a research programme on the dynamics of krill swarms- the programme should include routine plankton sampling from exploratory and commercial trawlers at each trawling station, collation of echo traces of krill swarms, and the deployment of drifting sensors in satellite communication; and that liaison be established with the working party co-ordinating underway observations from supply ships so that opportunities for time-series studies at the ice edge and in the pack ice from icebreakers and support ships are fully utilized. 3. That interest be invited and promoted in the development of a satellite communication drifting buoy assembly for studying the dynamics of krill swarms using sensors to measure zooplankton and phytoplankton biomass in the water column. 3.3 Shore-based studies So far only a few experiments have been successful on krill and other Antarctic plankton organisms in captivity on board research vessels and at shore stations. Basic data on productivity, respiration, growth, and reproduction, should be obtained together with data on rates of filtration, feedmg, digestion, and on metaboUsm under different controlled environmental conditions in tanks and aquaria. Attempts to culture krill experimentally should provide further information on the duration of various life history stages and on the biochemistry and histology of moulting and sexual maturation. Pressure tanks might be needed for the eggs and eariy larvae. Research on enzyme kinetics and microbial activity is needed in conjunction with studies on the energy budget and decomposition processes in large experimental enclosures. Recently, considerable experience has been gathered from plankton experiments in various types of inshore and land-based enclosures. Studies in feeding and predation of krill under semi-natural and semi-controlled conditions might bridge the gap between the physiological laboratory studies and field observations. Experiments on shoaling, vertical migration and sonar target responses of marine organisms such as krill in enclosures would be complementary to field studies. Year-round studies of the Antarctic fauna and fiora are essential because most species have marked seasonal cycles. The shore stations-although limited in their seagoing facilities-provide opportunities for continuous year-round observations of plankton and benthos in near-shore communities, including their predation by fishes, seabirds, and seals. The great number of experimental studies and their methodological diversity require considerable international and interdisciplinary collaboration in more than one well equipped shore station with easy access to krill stocks. Further information is required on the programmes, technical facilities and manpower of the various stations, in order to launch new internationally co-ordinated programmes as outlined above. 3.3.1 Role of established shore stations The current phase of Antarctic near-shore marine research was initiated during the International Geophysical Year. This near-shore research was made possible by the estabhshment of a number of permanent research stations on the Antarctic continent and on island groups in the Southern Ocean which were manned throughout the year. The principal roles of the established coastal stations in the BIOMASS programme are likely to be: to complement and add year-round continuity to the offshore ship-based studies; to analyse samples taken by the research vessels but which can be most successfully studied at the shore stations; and to study organisms which spend parts of their life histories in near-shore waters. H-55 PRACTICAL IMPLEMENTATION OF THE RESEARCH PROGRAMME 3.3.2 Treatment of pelagic samples Preliminary treatment of the biological material collected is best carried out on board research vessels and then transferred to a sorting centre. However, analyses which must be made soon after collection, especially those which require precision weighting, are probably best done at an Antarctic shore station. The biochemistry of the key organisms in the offshore regions at different trophic levels has yet to be described in any detail. Analyses of body tissue components are required to show seasonal variations and changes at different life stages. 3.3.3 Near-shore hydrography The wide geographical distribution of coastal stations in the Southern Ocean facilitates year-round, simultaneous and standardized observations of the physical, chemical and biological parameters of coastal waters. These observations wiU complement data gathered during the offshore programme. The routine techniques should be simple and unsophisticated in order to ensure the maximum participation by nations engaged in research in the Antarctic. Measurements should be made at fixed stations covering a long time series, as far as possible throughout the year. They should be taken at defined intervals and at standard depths. The most important parameters to be measured are: temperature, salinity, basic nutrients, light penetration, chlorophyll concentration, and concentration of suspended inorganic material, but it is desirable to make observations of as many relevant parameters as possible. It is important to discover how measurements taken near the coast in such a programme might differ from the data collected by oceanographic vessels. Therefore, as far as possible it will be necessary to initiate projects to quantify the differences between neritic and offshore waters, to describe the interaction between the near-shore and offshore environments and to determine the magnitude or trend of any differences between them. 3.3.4 Ecological studies and experimental research at shore stations Research will centre on investigations of the growth, metabolism, biochemistry, behaviour and reproductive biology of plankton and nekton species. The important potential research opportunities are: 1 . Near-shore species. (a) Broad based energy budget modelling of the near-shore water column in order to identify the principal components, the main routes and rates of energy flow, and its potential contribution to the offshore ecosystem. (b) Research on the trophodynamics of large concentrations of breeding seals and birds and their impact on the neritic and offshore ecosystems. (c) Research on the biology of species which have part of their life cycle in near-shore waters but also comprise potential offshore living resources (eg, Notothenia rossii). (d) Research on accessible neritic species which are closely related to important offshore species (eg, the neritic euphausiid, Euphausia crystallorophias). 2. Offshore species. (a) Research on offshore species such as Euphausia superba held in aquaria at shore bases, including experiments on metabolism, growth, feeding and food assimi- lation rates. (b) The provision of cultures of offshore phytoplankton for basic studies on the H-56 PRACTICAL IMPLEMENTATION OF THE RESEARCH PROGRAMME biochemistry and physiology of individual species and for feeding experiments with herbivores. (c) Analysis of the characteristics of the bioluminescence emitted by offshore species in connection with the proposals to use remote sensing of zooplankton (Section 2.10). (d) Controlled laboratory experiments to analyse the behaviour of appropriate offshore species, including shoaling and vertical migration, and to identify seasonal changes and changes during the ontogeny of each species. (e) Research on shoaling, vertical migration and sonar target responses of marine organisms such as krill, using plastic enclosures at sites such as Cumberland Bay, South Georgia. (f) Research on enzyme kinetics and micro-organism activity in conjunction with studies on the energy budget and decomposition processes. 3.3.5 Existing facilities There are 39 coastal stations operated by 10 countries. Of these, 14 are engaged in marine biological programmes and 1 1 in marine vertebrate studies. The following information is required: 1. Number and location of coastal Antarctic research stations and the months during which each base is operational. 2. Capacity of station, number of scientists and technicians and opportunities for visiting scientists. 3. Whether marine projects are undertaken, and the routine marine sampling procedures undertaken. 4. Period of year during which marine projects are undertaken. 5. Logistic support and facilities available (including number and size of boats, diving facilities, submersibles, aquaria, chemical laboratories, etc). 6. Details of capital equipment normally available for laboratory and field studies in marine biology, physiology and biochemistry. 7. Programme of current marine studies and those planned for the future. SELECTED REFERENCES Beamish, P. 1971. Quantitative measurements of acoustic scattering zooplankton organisms. Deep Sea Research, Vol 18, No 8, p 811-22. CreSWELL, G. R- 1976. A drifting buoy tracked by satellite in the Tasman Sea. Australian Journal of Marine Freshwater Research, Vol 27, p 25 1 -62. 3.4 Data reporting and handling 3.4.1 General The proposed large scale programme will generate information and data relating to all aspects of the living resources and their environment. Arrangements need to be made to ensure that individual scientists can easily locate all information (from whatever source) relevant to H-57 PRACTICAL IMPLEMENTATION OF THE RESEARCH PROGRAMME their particular studies. This will require some central organization (or organizations) which will have two rather distinct functions: (a) awareness, or cataloguing, and (b) information compilation and processing. The nature of the data handling problem also depends on the kind of data. For example, information on surface temperature or on the weight of a given species of fish caught in a commercial fishery, which can easily be expressed as single figures in standard and comparable units, presents different problems from complex biological information. Data from research vessels engaged in scientific work are always easier to obtain than data from commercial or semi-commercial operations. The mechanisms required are also likely to vary because different national agencies may be involved. The following sections discuss, first, the actual technical work needed with respect to data of different types and from different sources, and secondly, the facilities required to do this work, and the commitments that must be undertaken by countries participating in research or harvesting activities in the Southern Ocean. 3.4.2 Nature of activities 1 . Oceanographic data. These include all types of observations on the physical and chemical characteristics of the water masses, which are important for understanding the environment of the living resources; a study oF the impact of the physical environment on the abundance and distribution of these resources is especially important. Oceanographers, through national and world data centres, have well established arrangements for the exchange and central compilation of such data. Probably no new action needs to be taken other than to ensure that existing arrangements fully cover data from the Southern Ocean, with any minor modifications that seem desirable. Reporting, from whatever source, would probably be mainly the concern of IOC's co-operative programme in the Southern Ocean. 2. Catalogues of biological data. Biological information includes a wide variety of types, from data on sizes of animals in commercial catches (which would probably be best handled in the same way as general statistical data) to samples of plankton, which may need special co-operative arrangements for sorting and taxonomic identification. Between these extremes lie most biological observations (egy gross catches of major elements in plankton nets, observations on occurrence of echo-traces). Many of these observations are not suited for routine automatic processing because interpretation of the results {eg, the volume of the plankton net catch or the intensity of echoes) depends on quahtative information about the precise type of equipment, the way it was used and the type of organisms concerned. The major need with respect to these data is a register or catalogue, so that the interested scientist can readily discover what observations have been made and which organization or individual scientist should be approached for further information. 3. Sorting of biological samples. Many biological samples, particularly plankton and benthos samples, but to a certain extent catches of larger crustaceans, molluscs and fish, contain a variety of taxonomic groups whose detailed identification requires specialized knowledge. A large part of the information concerning the sample (eg, total weight of plankton in the haul or the number of copepods) may be obtained at the time of observation, or soon afterwards. However, the scientist or institution making the collection may not have the detailed knowledge of some or all of the taxonomic groups to enable them to complete the sorting and identification. This is best carried out in specialized centres to which samples can be sent when the gross analysis has been completed and more detailed information is required. Bearing in mind the lack of adequate knowledge of most of the living resources in the region, such procedure will involve a fairly H-58 PRACTICAL IMPLEMENTATION OF THE RESEARCH PROGRAMME significant sub-sample of the total observations, including catches of fish and other organisms by commercial or semi-commercial vessels. 4. Statistical and related data. It is convenient to consider together all of the biological and related data of a routine or semi-routine nature, particularly, but not exclusively, those arising from commercial or pilot scale observations. Such data include: statistics of total harvest (with details of species caught, and the location and time of capture); statistics of fishing effort (types of vessel and gear used, number of hauls, days of operations, etc). The new data from individual countries or expeditions need to be combined to provide best estimates of quantities such as: the total catch of a species in a certain area during a particular period; the seasonal and year-to-year changes in abundance and distribution, as measured by changes in catch per unit effort; changes in size composition of the catches. The need to check on the reliability, consistency and comparabihty of the data can be time consuming; much of the information will come from commercial sources which may have little immediate interest in the quality of the data submitted. Again, those actively involved in harvesting a resource, but who have not submitted information, will need to be contacted. The data then have to be combined and published in tabulated forms (statistical bulletins, sampling reports) similar to those issued by regional fishery bodies in other parts of the world. This is presently done by the Bureau of International Whaling Statistics with respect to whaling; however, the volume of work would be much greater if harvesting of other resources began on a large scale. 5. Bibliographic information. As the volume of publications on the living resources of the Southern Ocean grows, it becomes more difficult for the individual scientist without assistance to become aware of all significant new publications and to identify relevant material in the past literature. At least three information systems-the Cold Regions Bibliography Project, Library of Congress (which pubUshes Current Antarctic Literature and the Antarctic Bibliography); the Aquatic Sciences and Fisheries Information System, sponsored by FAO and other agencies (which publishes Aquatic Sciences and Fisheries Abstracts); and Recent Polar Literature, published by the Scott Polar Research Institute-are concerned with the living resources of the Southern Ocean, although they approach the subject as part of three quite different larger themes. There would seem to be little problem for these information services to produce specialized bibliographies of the so-called conventional hterature (ie books, papers in journals) covering the interests of a SCAR project. There would be advantages in this being done by all three (preferably in conjunction) in view of their different coverage of technical and scientific journals. The non-conventional literature, including mimeographed reports for meetings, and working party reports, is becoming increasingly important. In particular, most of the up-to-date information on exploratory and experimental fishing for. krill is contained in this type of literature. Since such information is not always published in the normal scientific hterature without long delay, it is essential that every effort is made to ensure wide distribution of the 'unconventional' literature and that information on availability is provided to appropriate centres. 3.4.3 Facilities and commitments required 1. General. None of the information systems briefly outlined above will work unless proper facilities are available, and unless those individuals or institutions with the information provide it to the relevant system. Much of the work could be carried out within existing arrangements (though possibly some minor additional funding would be required to do the work effectively), while some would need some essentially new arrangements. H-59 PRACTICAL IMPLEMENTATION OF THE RESEARCH PROGRAMME Oceanographic data can presumably be handled by existing national and world data centres; the commitment to supply data to these centres is generally accepted by oceanographers. Catalogues of biological data could also (in principle) be handled in the same way, though the volume of work may require additional funding. The existing ROSCOP and ROMBI forms may need adjustment for the particular interests of the Southern Ocean studies and all biological investigators will have to agree to complete these or some other type of report. This should present no problem with respect to observations made in the course of purely scientific studies, but may be more difficult in commercial or pilot scale operations. 2. Sorting of biological data. Since study of the biological environment of the Southern Ocean will often require processing and identification of specimens by persons not involved in the actual collection, it is important that each sample be uniquely identified at the time of collection and throughout later examination and identification. Standardized data recording and processing techniques were adopted on USNS Eltanin, which operated under the United States Antarctic Research Program (USARP) during its later cruises. Data recording forms (of which there were different kinds for pelagic and benthic sampling) were provided to all participants. Each form, in triplicate, was uniquely numbered; this number was henceforth the identification of the samples and the reference to all relevant data. In addition to recording basic data, forms for the study of living resources were also designed to record observations on abundance, size of organism, etc. These procedures accomplished at least two important objectives. (a) There was no longer any confusion caused by different investigators assigning the same series of numbers to entirely different samples. (b) Availability of forms, with estabUshed data fields, encouraged participants to record more observations in a consistent manner-a particularly relevant objective for large programmes in which the collecting operations wiU involve personnel who are less experienced than the programme directors. Experience has shown that the sampling data should be summarized and reviewed for errors prior to the end of the cruise or soon thereafter. Recording of data on multi-copy forms facilitates this procedure of data quality control. The Smithsonian Oceanographic Sorting Center (SOSC) has developed a computerized biological station data programme for data exchange and analysis. Data may be entered into a generalized master file that can be augmented as new information (such as identifications or bibliographic references) becomes available. With such a file, data and standard or special reports are readily available to the investigators. In addition, the file facilitates the publication and distribution of technical data in a timely and reliable manner. Participating countries can acquire the computer system and process data directly or send information to a centralized location for processing and distribution to all participating countries and individual scientists. 3. Statistical and related data. As noted above, this work of data collection would be quite extensive once any significant harvesting (other than whales) began. Regional fishery commissions usually need several people (including clerical staff) to handle their statistical work. Technically, several institutions, including FAO, could carry out the work but would need additional funding. (FAO already handles these data on a global scale but without detailed breakdown except in those regions for which FAO has special responsibility. The lack of detail causes difficulty in interpretation; for example, the region of harvest is given in no more detail than, say, the south-west Atlantic, which covers an area from South Georgia to northern Brazil.) A more serious problem might be that of ensuring fully detailed collection and reporting of H-60 PRACTICAL IMPLEMENTATION OF THE RESEARCH PROGRAMME data from commercial operations. Countries becoming members of regional fishery commissions normally accept commitments to supply data to the commission; a similar arrangement (in this case, to report data to SCAR and to other contracting parties) is contained in Article 5 of the Convention for the Conservation of Antarctic Seals. Arrangements and commitments of the same kind may need to be made with respect to any harvest made in the Southern Ocean. 3.4.4 Proposak and recommendations Arrangements for exchange of information, and for the reporting, compilation and dissemination of appropriate data will be important features of the proposed programme. While many details can only be settled when the extent of research programmes and the scale of commercial exploitation are better known, the following specific actions can be recommended. 1 . All oceanographic data should be reported to relevant centres. 2. Statistics of commercial and exploratory fishing should be reported to FAO along the lines set out in Appendix A, using the standard global classification of area and species. The proposed northward movement of the boundary lines between statistical areas in the Atlantic and Indian oceans was agreed to, and in the interim period before the new regions are formally approved by all interested parties, countries should distinguish separately, when reporting to FAO, the catches taken: (a) in the South Atlantic in the area bounded by 50° to 60°S in 20° to 50°W and 55° to 60°S in 50° to 60°W, and (b) in the Indian Ocean between 45° and 50°S in 30° to 80°E. Countries should also distinguish the major species as listed in the relevant section of this proposal. In view of the importance of proper statistics, it is recommended to SCAR that it draw the attention of Antarctic Treaty countries and other interested parties to the reporting of data on the catches taken. 3. Consideration should be given to the needs and possible arrangements for the exchange and storage of biological data, other than fishery data which could be handled by FAO. 4. Countries are urged to exchange early information on research being planned and being undertaken. H-61 4. INTERNATIONAL CO-ORDINATION AND CO-OPERATION There exists a number of international organizations, at various levels, which have expressed interest in the resources of the Southern Ocean (see Figure 6); several of these have active biological programmes. The success of BIOMASS will depend on the establishment of effective co-ordination machinery. MEMBER COUNTRIES Non-Qovernmen IOC MEMBER STATES IINTERESTEDI gl orggnizcrtions: Govemmentgl orqgnizalion5 ICSU lABO SCOR SCAR ACMRR TREATY SCOR/SCAR ' — "^ Group of Speciglists IOC lUNESCOl ICG SouthGfn Ocegn [ICG BIOMASS Sub-group — > Line of commgnd -■^ Advice c^ IWC Scientific pjgnning and ewluotion y Co-ordingtion of operations KIG 6. Organizational structure for BIOMASS. 4.1 Present organizations 4. 1 . 1 Non-govemmental organizations 1. ICSU-SCAR. 2. ICSU-SCOR. 3. ICSU-IABO. 4.1.2 Inter-governmental organizations 1. Antarctic Treaty consultative meetings. 2. Biennial meetings of representatives of consultative governments. H-62 INTERNATIONAL CO-ORDINATION AND CO-OPERATION 3. Intergovernmental Oceanographic Commission (IOC) of UNESCO. (a) While the IOC is constitutionally attached to UNESCO, it serves, with respect to marine research, as a 'joint specialized mechanism' of other interested UN bodies, specifically, FAO, WMO and IMCO, as well as the UN itself (ICSPRO organizations). (b) IOC has a subsidiary body, an International Co-ordination Group for the Southern Ocean (ICG/SOC), established in 1970. It is composed of IOC member states interested in Antarctic research, with observers from SCOR, SCAR, ACMRR (FAO) and other interested organizations. The group is concerned with all aspects of Southern Ocean scientific studies and will need to maintain close co-ordination between BIOMASS and other SOC activities. ICG/SOC has the present terms of reference: (i) To assemble and distribute details of firm oceanographic cruise plans in the Southern Ocean, preferably at least one year in advance, (ii) To encourage the pre-allocation of blocks of time for oceanographic research on Antarctic supply vessels whenever practicable, (iii) To develop means of co-ordinating existing and planned oceanographic research programmes in the region, (iv) To encourage the evaluation of existing oceanographic data from the region with the intention of fostering specific studies of Umited extent and capable of being carried out in the foreseeable future, (v) To encourage and review the development of relevant theory, methods and instruments with particular reference to the problems of obtaining measurements in the winter and in the presence of ice. (vi) To develop plans for the gradual evolution of a comprehensive study of the Southern Ocean. (c) Having considered the recommendations of the second session of ICG/SOC (Buenos Aires, 15-19 July 1974) the IOC Executive Council invited the SCAR Group of Specialists on Living Resources of the Southern Ocean to prepare practical proposals for collaborative investigations on the biological oceanography of the area and for the organization of multi-ship studies. (d) Global Investigation of Pollution in the Marine Environment (GIPME): the intergovernmental Oceanographic Commission has published 'A comprehensive plan for the global investigations of pollution in the marine environment and basehne study guidelines' (IOC Technical Series No 14). Any activities in the Southern Ocean promoted under this programme by the IOC Working Committee for GIPME should be co-ordinated with BIOMASS. 4. The Food and Agriculture Organization of the UN (FAO). The FAO has direct interest in the resources of the Southern Ocean and is already active in several spheres. Its main interests are in the better utilization and management of these resources rather than in scientific research in the narrow sense. However, a number of FAO activities are directly relevant to the BIOMASS project. (a) The ACMRR of FAO (a body of scientists which, like SCOR, is also advisory to IOC) is presently active in evaluating the status of Antarctic resources of seals and cetaceans. The study is financed by UNEP, bUateral funds, and by FAO itself, and culminated in the World Scientific Conference on Marine Mammals held in Bergen in September 1976. An important feature of that conference was the formulation of a long term research programme on marine mammals. There will need to be close co-ordination between the proposed research programme concerning Antarctic stocks of seals and cetaceans and BIOMASS. (b) FAO in 1974 convened an 'Informal consultation on Antarctic krill' (published as FAO/Fisheries Report, No 153). At this consultation the FAO Department of Fisheries agreed to: H-63 INTERNATIONAL CO-ORDINATION AND CO-OPERATION (i) Produce a bibliography on Antarctic krill. (ii) Produce updated accounts of the knowledge of the resource, its exploitation and utilization, (iii) Act as an information centre on research plans, resource knowledge, fishing techniques, equipment and utilization questions, (iv) Review (when appropriate) the need for international action such as ad hoc expert group meetings and joint research projects, and pursue the possibility of mobilizing international financing for resources survey work and other activities of a high priority. Preliminary drafts of the bibUography and of the resource reviews, and an account of the exploitation and utilization of krill were presented to the Woods Hole meeting. (c) FAO is the executing agency for a proposed Southern Ocean Fisheries Programme to be funded by the United Nations Development Programme. The preliminary phase of this programme started in July 1975 and is helping to support, among other activities, the resources reviews referred to in Section 2. The programme's main phase will include the compilation and dissemination of resource information. In addition, surveys of potentially exploitable resources, especially those which complement activities of other institutions, will be made to the extent of available support from UNDP and other funding agencies. (d) FAO compiles and publishes statistical information on commercial fish catches and species categories by broad areas and on a worldwide basis. These data can provide essential basic information for resource studies, particularly if the regional and species classifications of Antarctic and sub-Antarctic catches were improved. 5. The International Whaling Commission (IWC). The IWC has launched an International Decade of Cetacean Research, in response to proposals from UNEP and others. It has asked UNEP, FAO and perhaps other bodies for funds. It has been suggested that the World Bank might also be interested. IWC has also just embarked on a 'new management policy' for whaling by its member states, which requires considerably expanded research for effective application. The policy should reduce uncertainties in whale stock assessment and resolve problems arising from interactions between species. 6. United Nations Environment Programme (UNEP). UNEP has a wide interest in environmental affairs, including the conservation of natural resources. It has helped support the ACMRR (FAO) study of marine mammals mentioned above. 7. International Union for the Conservation of Nature and Natural Resources (lUCN). Quasi inter-governmental, lUCN has several levels of interest in Southern Ocean research: they range from expressions by its General Assembly of a more or less political nature, as to the desirable status and uses, or non-uses, of tlie Antarctic and the surrounding seas, to the various specialized groups— on whales, seals and now on marine mammals as a whole-under its Survival Service Commission. lUCN will be concentrating in 1977-79 on marine activities. H-64 5. PROVISION FOR SCIENTIFIC ADVICE 5.1 Introduction A major objective of the BIOMASS programme is to help ensure the wise and careful conservation and utilization of the hving resources for the benefit of present and future generations of mankind. In particular, we hope to avoid the mistakes that have caused the collapse of many fish stocks. In view of these mistakes, it should be reaUzed that the BIOMASS programme is a vital first step towards providing the necessary scientific information on different resources and their interactions. However, this will not ensure the wise use of these resources. Two other steps need to be taken: (a) examination of the technical aspects of different management strategies; and (b) presentation of the results of the scientific studies in a manner that will facilitate decision making by the governments concerned. 5.2 Examination of management strategies Management, even of a single species, can be carried out in a number of different ways, and the number of alternatives increases rapidly when management is involved with a number of distinct but interacting resources. Management considerations in the Southern Ocean should include investigations on the economics and social condition of the fishery, as well as on its biology, since these can be critical in determining the success or failure of a management strategy. For example, one important weakness of the initial regime established by the IWC was that the overall Antarctic quota was not allocated to countries or to individual enterprises. As a result, a considerable over-capacity developed in the fishery during the I950's; by 1964 the industry was in poor economic shape and not in a position to accept readily the substantial cuts in quota that the IWC Committee of Three then deemed to be necessary. Conmiercial scale harvesting of krill may start in the near future, and it is unlikely that firm and precise estimates of the potential annual yield, (ie, the quantity that can be taken year after year without serious damage to the stocks of krill or other species) will be available by then. Two extreme reactions to this situation are: (a) to allow completely unrestricted fisheries for krill untU the need for specific measures becomes obvious, or (b) to ban harvesting untU better data are obtained. The dangers of the former, which has been the strategy impHcitly adopted for many resources in the past, are now well known. The disadvantages of the latter are also great; first, it sacrifices the benefits that could accrue from adequately controlled krill harvesting during the interim period; second, it would be difficult, if not impossible, to determine the potential sustained harvest until some harvesting is done. Without significant exploitation, it is also difficult to estimate the effects of different levels of harvest on the detailed characteristics of the krill populations' abundance, growth, mortality or reproductive rates and on the other populations (whales, birds, etc) that interact with krill. The preferable strategy is one that allows some degree of harvesting, but under careful control and with arrangements to ensure that the harvest can be reduced quickly if there is any evidence that quotas have been set too high. It would, of course, be easier to detect changes if the harvest were concentrated in some limited area. The information obtained on the effects of harvesting on ecosystem relationships could then be used to develop management strategies for the whole area. This plan seems appropriate even though the population boundaries are still poorly defined. The required studies will involve a variety of expertise in such subjects as economics, and biology (especially population dynamics). The model described in Section 2.2 of this proposal H-65 PROVISION FOR SCIENTIFIC ADVICE will obviously be basic but will need to be supplemented by careful analytical studies by experts in the fields concerned. The basic studies of individual stocks, proposed in Section 2, will be vital parts of the exercise. 5.3 Arrangements for providing advice Administrators and other decision makers need scientific advice in a readily understandable form. Also, the management process, based in part on the scientific evidence, needs to be kept quite separate from the process of reaching agreement among scientists on interpretation of the scientific evidence. Experience in many fishery commissions has shown that progress towards administrative or political agreement on particular measures can be seriously impeded if scientific arguments are re-opened during the discussion. For example, negotiations on how to limit the catch of certain species to 100 000 tons, involving possible severe constraints on at least some fishermen, will soon break down if it is suggested that perhaps the limit need not be lower than 150 000 tons. Agreement between scientists in this sense does not necessarily mean unanimous agreement on some single figure. It must be expected, given our uncertainty about the structure and functioning of the Southern Ocean ecosystem, that there will be considerable differences in interpretation. Any scientific statement or advice used in management decisions must take these doubts and differences into account. This can be done within a single agreed report, provided the differences and their implications (especially with regard to the results of different management measures) are spelt out. If such a report is to be accepted and used by the decision makers as a single agreed document, all concerned scientists must, in principle, be able to participate in the discussions and the preparation of the report. In fishery commissions this function is usually carried out by a 'standing committee on research and statistics' (a title that reflects the importance of statistics and similar data to the whole process) or an equivalent body set up within the commission. However, in the case of the two commissions in the north-east Atlantic (including the Baltic) this function is carried out by the International Council for the Exploration of the Sea. For practical reasons, the main work is often done by small subcommittees or working groups, usually consisting of specialists on stock assessments. The experience of the fishery scientists working in these groups is that groups of specialists, though normally nominated by and representative of their national governments, do work effectively together and are generally highly successful in carrying out the scientific work. The degree of co-operation and success tends to increase with time, particularly if the group has had the chance to work together before difficult and politically sensitive problems of management occur. The fishery commissions often have other standing committees or groups to examine technical, but wholly biological, questions concerning economic, social or other problems related to determining alternative management measures. Similar arrangements for detailed examination and review of scientific and technical aspects may be desirable for the Southern Ocean. Although consideration of the structure of such a group is outside the scope of this document, once large scale exploitation starts and the responsibilities of the scientific planning group increase, its effective functioning will require some semi-permanent secretariat to provide support regarding such matters as compiUng routine data, processing reports, and servicing meetings (which are likely to become fairly frequent). It may be desirable to arrange for additional contributions to the group from those experts who are concerned with more long term interests, particularly in relation to the problems of the world community as a whole. The introduction and implementation of management measures will require some formal inter-governmental arrangements; the estabUshment of the necessary structure will call for extensive international discussions, taking into account a large number of factors, of which scientific research will be only one. H-66 TIME-TABLE FOR BIOMASS The three major types of activity in the BIOMASS programme (seagoing experiments and surveys; shore-based experiments and year-round observations; data analysis and modelUng) have to be adjusted to each other. Observations on krill swarms and on the ecological physiology of key organisms of the Antarctic ecosystem should be started immediately and continued both at sea and at shore stations. Systems of data reporting and handUng should be developed as soon as possible. The programme by FAO as discussed and approved by its Committee on Fisheries includes the development of a statistical data base and of a collection of scientific reviews on the Uving resources and their present and potential exploitation. Some standardization with regard to sampling and sorting of plankton and nekton in the Southern Ocean is also required at an early stage of the BIOMASS programme. 1977 Saenlrfic ptonnng . 1978 1979 1980 1981 1982 1983 1984 1985 1986 ^^^^^^^^ TecnfKOl prepofotion L _^^| Seogoing experiments and surveys FIBEX SIBEX Other seogbrg octivities 1 m-M ^^ Share based stucies ^m ^ ■■ ^m ^Jh_^h Oato analysis , SynthesB and advice, (workshops and symposia 1 ■ 1 J 1 FIG 7. Time-table for BIOMASS. The main implementation phase consists of at least two major international multi-ship experiments as well as of a number of loosely connected experiments at sea and ashore and large scale surveys at the national level. A time-table for large scale seagoing operations has not only to take into account other major oceanographic activities such as FGGE but also the need for sufficient time for scientific planning, logistic preparation, and development of instrumentation. The austral summer 1980-81 has been chosen for the First International BIOMASS Experiment (FIBEX)-a major multidisciplinary process-oriented study involving several vessels in small scale experiments and large scale investigations of transport mechanisms and ecosystem H-67 TIME-TABLE FOR BIOMASS variability in space and time. Operations will concentrate on the Atlantic sector but include Drake Passage and its western approaches. Current marine research activities in the Antarctic should be used as far as possible to prepare the ground for this first BIOMASS experiment. Data analysis will be carried out in conjunction with field work, laboratory studies and modelling. Small groups of specialists will need to meet to plan seagoing and shore-based experiments, and to develop methodology, data handling and modelling concepts. An interim synthesis of data and results should be presented at specialized workshops from 1979 onwards and at a small symposium in 1982. The complete results should be ready for presentation at a major symposium in about 1986. H-68 7. PLANNING AND CO-ORDINATING GROUPS Preparatory and co-ordinating arrangements for the implementation of BIOMASS will be needed at the scientific and administrative, non-governmental and governmental levels. The group felt that such arrangements should largely be entrusted to the existing bodies of SCAR/SCOR and of IOC, which should also be the principal sponsors of BIOMASS. The group proposed that a SCAR/SCOR scientific planning group be established which would recommend methods and techniques and draw up scientific programmes for co-operative field work and shore-based studies and for the analysis of data. Sub-groups could be established for special technical tasks. The existing Group of Specialists on the Living Resources of the Southern Ocean (SCOR WG 54) could, with perhaps enlarged membership and its new terms of reference, undertake such a task. The group further proposed the formation of a BIOMASS co-ordinating group as a sub-group of the existing IOC International Co-ordination Group for tlie Southern Ocean (ICG/SOC), with responsibility for the implementation of the BIOMASS programme acting on the scientific advice of the SCAR/SCOR scientific planning group. Consideration might be given to the appointment of an international co-ordinator. SCAR. SCOR and lABO should be invited to participate in the work of the BIOMASS co-ordinating group, and it is expected that FAO and the International Whaling Commission will contribute to the efforts of this group. The proposals on the co-ordination of BIOMASS and on exchange of information and data are summarized in the recommendations given in the following section. H-69 8. COMPILATION OF RECOMMENDATIONS ON THE ORGANIZATION OF BIOMASS The Group recommends: 1. That SCAR and SCOR approve tJie following amended terms of reference for the Group of Specialists on Living Resources of the Southern Ocean (SCOR WG 54). (a) To encourage and stimulate investigations of the trophodynamics of the Antarctic marine ecosystem and the ecology and population dynamics of organisms at different trophic levels. (b) To keep under review the current state of knowledge concerning the Antarctic marine ecosystem from the viewpoint of structure, biomass and dynamic processes of organisms at different trophic levels. (c) To transmit through SCAR and SCOR to those countries engaged in the exploitation of the living resources of the Southern Ocean scientific information relevant to the management of such resources. (d) To advise SCAR and SCOR and through them other international organizations, and in particular to respond to relevant recommendations of IOC and the Antarctic Treaty consultative meetings. (e) To act as the international scientific planning group for BIOMASS. (f} To recommend standardized measurements, techniques and data recording of biological observations in the Southern Ocean. 2. That IOC undertake the international co-ordination of BIOMASS. 3. That IOC request countries carrying out research in the Southern Ocean to provide details of proposed cruise tracks and scheduled researches, which would be made available to the Group of Specialists. 4. That SCAR request the National Agencies operating supply sliips to institute a circum-Antarctic programme of underway observations of surface temperature, salinity, chlorophyll and underway collection of records of expendable bathermograph (XBT) and biological echo traces. 5. That SCAR collaborate with FAO in drawing the attention of all parties engaged in the exploration and exploitation of living resources of the Southern Ocean to the need for detailed catch and effort statistics as set out in Annex A to be submitted to FAO. 6. That SCAR inform FAO of its approval of the proposed northward movement of the boundary Hues between statistical areas in the Atlantic and Indian oceans, and that in the interim period before the new regions are formally approved by all interested parties, countries should be requested to distinguish separately, when reporting to FAO, the catches taken (a) in the south Atlantic in the area bounded by 50° to 60°S in 20° to 50°W and 55° to 60°S in 50° to 60°W, and (b) in the Indian Ocean between 45° and 50°S in 30° to 80°F. H-70 APPENDIX A Statistics from commercial fisheries As discussed in the main proposal (particularly Section 3.4.3), it is essential for the success of BIOMASS, and for the rational utilization and management of the resources, that scientists have available to them detailed statistics from any commercial fisheries in the Antarctic. The following proposals have been made regarding the details of the statistics that should be reported to FAO as soon as possible following the end of each Antarctic season: 1 . The catches. The quantities should refer to nominal catches, ie the live weight of the animals as they leave the water, before any processing takes place. These nominal catch data, reported in metric tons, should be broken down as follows: (a) Time periods: These should be in calendar months. Annual summaries should be for split years (1 July-30 June). (b) Species items: These categories should refer preferably to individual species, but in some cases (for example, cephalopods) to groupings by genera or even families. (c) Fishing areas: The catch data reported should refer to quadrangles of 5° latitude by 5° longitude; if this amount of detail cannot be provided, then the areas should refer to three FAO 'Major Fishing Areas' for statistical purposes, ie FAO Areas 48, 58 and 88. (d) Fishing gear: The catches should be broken down by the principal fishing methods used (for example, trawls, purse-seines, longlines, etc) listed in FAO's International Standard Statistical Classification of Fishing Gear. 2. Fishing effort and corresponding catches. For nominal catches (reflecting the details discussed in the preceding paragraph) the corresponding fishing effort should also be provided for each 'kind of fishing unit'. The fishing unit is determined primarily by the gear used, and secondarily by the size (CRT) of the vessel. The information on fishing effort should cover 'fishing time' and 'fishing power'. 'Fishing time' concepts are related to the type of gear used. For trawls, the principal effort measure is 'number of hours fished', for purse-seines the 'number of sets'. Standard definitions for these and other effort concepts have been established by the Co-ordinating Working Party on Atlantic Fishery Statistics and are under constant refinement and review by that body. 3. Fishing fleets. Details should be given by national offices on all the individual units (mother ships, factory vessels, fishing vessels, etc) of their fleets operating in one or more of the three FAO Antarctic Fishing Areas during a specified split year, 1 July-30 June. For each of these vessels such information should be provided as: (a) its nationality and name, (b) type of vessel, (c) overall length, (d) gross registered tonnage, (e) horsepower, (0 type or types of fishing gear, (g) person- nel, (h) hold capacity for various products, (i) special processing equipment, electronic equipment, etc. The following documents, available from the Fishery Statistics Unit, Department of Fisheries, FAO, Rome, provide detailed information on statistical standards and requirements: A proposed international statistical system for the Antarctic fisheries-FAO Fishing Areas 48, 58 and 88 (FAO Fisheries Circular No 608). World chart: Major fishing areas for statistical purposes (FAO Fisheries Circular No 420). H-71 APPENDIX A: 'Nominal catches' and 'Landings': Definitions and notes (FAO Fisheries Circular No 428). Notes on international classifications and definitions used in fishing fleet, fishing gear, fishing effort and fishermen statistics (FAO Fisheries Circular No 429). «-72 APPENDIX B Participants SCAR/SCOR Conference on Living Resources of the Southern Ocean Academy Summer Study Center Woods Hole, Massachusetts, USA 17-21 August 1976 Dr David G. Ainley, Point Reyes Bird Observatory, Mesa Road, PO Box 321, Bolinas, California 94924, USA. Dr Vera Alexander, Institute of Marine Science, University of ,Maska, Fairbanks, Alaska 99701, USA. Dr Patrick M. Amaud, Station Marine d'Endoume, rue de la Batterie des lions, 13007 Marseille, France. Mr Arthur de C. Baker, Institute of Oceanographic Sciences, Brook Road, Wormley, Godalming, Surrey 9U8 SUB, England. Dr Gerald Bertrand, Council of Environmental Quality, Executive Office of the President, 722 Jackson Place NW, Washington DC 20006, USA. Mr W. Nigel Bonner, Life Sciences Division, British Antarctic Survey, Madingley Road, Cambridge CB3 OET, England. Dr Robert L. Brownell, Division of Mammals NHB-398, National Museum of Natural History, Smithsonian Institution, Washington DC 20560, USA. Dr P. R. Condy, Mammal Research Institute, Department of Zoology, University of Pretoria, Pretoria 0002, South Africa. Dr David L. Cram, Sea Fisheries Branch, Department of Industries, Private Bag, Seapoint 8060, Cape Town, South Africa. *Sir George Deacon, Institute of Oceanographic Sciences, Wormley, Nr Godalming, Surrey 9U8 SUB, England. Dr John H. Dearborn, Department of Zoology, University of Maine, Orono, Maine 04473, USA. Dr Robert de la Fortelle, SAPMER, 32 rue La Boetie, 75016 Paris, France. Dr Arthur L. DeVries, Scripps Institution of Oceanography, University of California, San Diego, La JoUa, California 92037, USA. Dr Hugh H. DeWitt, DarUng Center for Research Teaching and Science, University of Maine, Walpole, Maine 04S73, USA. Dr Thang Do Chi, Universite des Sciences et Techniques de Languedoc, Laboratoire d'Hydrobiologies, Place Eugene Bataillon, 3400 MontpelUer Cedex, France. *Dr Sayed Z. El-Sayed {Convenor), Department of Oceanography, Texas A and M University, College Station, Texas 77843, USA. Dr Inigo Everson, Food and Agriculture Organization of the UN, Via delle Terme di Caracalla, 00100 Rome, Italy. Dr Theodore Foster, Scripps Institution of Oceanography, Marine Physical Laboratory, 132 Sverup Hall, La JoUa, California 92037, USA. •Members of SCAR/SCOR Group of Specialists on Living Resources of the Southern Ocean. H-73 APPENDIX B Dr Robert A. Frosch, Applied Oceanography, Woods Hole Oceanographic Institution, Woods Hole, Massachusetts 02543, USA. Dr Katherine A. Green, Biological Systems Modelling, 1 1801 Rockville Pike No 802, Rockville, Maryland 20852, USA. Dr George D. Grice, Jr, Department of Biology, Woods Hole Oceanographic Institution, Woods Hole, Massachusetts 02543, USA. *Dr John A. Gulland, Aquatic Resources and Survey and Evaluation Service, Department of Fisheries, Food and Agriculture Organization of the UN, Via delle Terme de Caracalla, 00100 Rome, Italy. *Professor Dr GotthilfHempel, Institut fiir Meereskunde an der Universitat, Diisternbrooker Weg 20, D23 Kiel, Federal Republic of Germany. Dr Robert J. Hoftnan, Marine Mammal Commission, 1625 Eye Street NW, Washington DC 20006, USA. Dr Osmond Holm-Hansen, Scripps Institution of Oceanography, Sverup Hall 2153, University of California, La Jolla, California 92037, USA. Dr Takao Hoshiai, National Institute of Polar Research, 9-10, Kaga 1-chome, Itabashi-ku, Tokyo 173, Japan. Dr Warren A. Hovis,Jr, FOB No 4, Room 135, Code S3 2, Washington DC 20233, USA. Dr Akito Kawamura, Whales Research Institute, 1—3-1 Etchuji*na, Koto-ku, Tokyo, Japan. *Professor George A. Knox, Department of Zoology, Private Bag, University of Canterbury, Christchurch 1 , New Zealand. Ms Betty J. Landrum, Smithsonian Oceanographic Sorting Center, Smithsonian Institution, Washington DC 20560, USA. *Dr Richard M. Laws, British Antarctic Survey, Madingley Road, Cambridge CB3 OET, England. Dr George A. Llano, Division of Polar Programs, National Science Foundation, 1800 G Street NW, Washington DC 20550, USA. Capitan Heman Lorca-Fuller, Instituto Antartico Chileno, Avenue Luis Thayer Ojeda 814, Correo Sucursal 21, Santiago, Chile. Dr C. C. Lu, Department of Biology, Memorial University of Newfoundland, St John's, Newfoundland, AlC 5S7, Canada. Professor Mary Alice McWhinnie, Department of Biological Sciences, De Paul University, 1036 Belden Avenue, Chicago, Illinois 60614, USA. Dr Olegl. Mamayev, IOC, UNESCO, 7 Place de Fontenoy, 75700 Paris, France. Licenciado Enrique Marschoff, Instituto Antartico Argentino, Cerrito 1248, Buenos Aires, Argentina. Dr Ole A. Mathisen, Fisheries Research Institute WH-10, College of Fisheries, University of Washington, Seattle, Washington 98195, USA. Dr Colin Nash, International Center for Living Aquatic Resource Management, Oceanic Institute, Waimanalo, Hawaii 96734, USA. Dr Keiji Nasu, Japan Marine Fisheries Resource Research Center, 3—4 KlO-1— chome, Chiyoda—ku, Tokyo 102, Japan. *Dr Takahisa Nemoto, Ocean Research Institute, University of Tokyo, Nakano, Tokyo 164, Japan. *Mcmbcrs of SCAR/SCOR Group of Specialists on Living Resources of the Southern Ocean. H-74 APPENDIX B *Dr Garth G. Newman, Sea Fisheries Board, Department of Industries, PO Box 251, Cape Town, South Africa 8000. Dr kvuzo Ohyama, Far Seas Fisheries Research Laboratory, 1000 Orido, Shimizu 424, Japan. *Dr Steinar Olsen, Institute of Fishery Technology Research, PO Box 1904, E. Sundsgt., 501 1 Bergen, 57 Nordnes, Norway. Dr Jan Piechura, Sea Fisheries Institute, Al. Zjeduocreme 1, Ckiynia, Poland. Dr T. Pommeranz, Institut fiir Meereskunde an der Universitat, Diisternbrooker Weg 20, D23 Kiel, Federal Republic of Germany. Professor Jean Prevost, UER des Sciences, Laboratoire de Biologie Animale, 123 rue Albert-Thomas, 87100 Limoges, France. Dr Stanislaw Rakusa-Suszczewski, Institute of Ecology, Polish Academy of Sciences, Dziekanow, Warsaw, Poland. Dr Ovde F. E. Roper, National Museum (Natural History), Smithsonian Institution, Washington DC 20560, USA. Dr Brian J. Rothschild, National Oceanic and Atmospheric Administration, National Marine Fisheries Service, Washington DC 20235, USA. Professor Dr Dietrich Sahrhage, Bundesforschungsanstalt fiir Fischerei, Palmaille 9, 50 Hambui^, Federal Republic of Germany. Dr Joachim Scharffe, Fish Production and Marketing Service, Fisheries Industries Division, FAO, Via delle Terme de Caracalla, 00100 Rome, Italy. *Dr Donald B. Siniff, Department of Ecology and Behavioural Biology, University of Minnesota, Minneapolis, Minnesota 55455, USA. Dr Finn Sollie, Fridtjof Nansen Foundation, Polhogda 9324, Lysaker, Norway. Dr H. Burr Stenbach, Oceanic Foundation, Makapuu Point, Waimanalo, Hawaii 96795, USA. *Dr Aldo P. Tomo, Instituto Antartico Argentino, Cerrito 1248, Buenos Aires, Argentina. *Dr David J. Tranter, Marine Ecosystems Group, CSIRO, Division of Fisheries and Oceanography, Post Office Box 21, Cronulla, New South Wales, Australia. Dr Jose Valencia, Instituto Antartico Chileno, Avenue Luis Thayer Ojeda 814, Correo Sucursal 21, Santiago, Chile. Dr John P. Wise, Resource Assessment Division (F17), National Marine Fisheries Service, Department of Commerce, 3360 Whitehaven Street NW, Washington DC 20235, USA. *Mcmbcrs ol SCAR/SCOR Group of Specialists on Living; Resources ol' the Soutlieni Oeeaii. H-75 ACMRR CARPAS FAO FGGE FIBEX GARP GIPME lABO ICSEAF ICSPRO ICSU IMCO IOC IOC's ICG for the Southern Ocean ISOS lUBS lUCN IWC NASA ROMBl ROSCOP SCAR SCOPE SCOR SOC UNDP UNEP UNESCO WMO APPENDIX C Acronyms Advisory Committee on Marine Resources Research of FAO Commission Assessores Regionale del Pesquerias del Allan tico Suroest (members: Brazil, Uruguay, Argentina). Subsidiary body of FAO, set up under Article VI of FAO constitution. Food and Agriculture Organization of the United Nations First GARP Global Experiment First International BIOMASS Experiment Global Atmospheric Research Programme Global Investigation of Pollution in the Marine Environment International Association for Biological Oceanography of lUBS International Commission for the Southeast Atlantic Fisheries (members: Spain, Portugal, Japan, USSR, Bulgaria, Poland, South Africa). Independent, H.Q. in Madrid. Inter-Secretarial Committee on Scientific Programmes Related to Oceanography International Council of Scientific Unions Intergovernmental Maritime Consultative Organization (UN) Intergovernmental Oceanographic Commission IOC's International Co-ordination Group for the Southern Ocean International Southern Ocean Studies International Union of Biological Sciences International Union for the Conservation of Nature and Natural Resources International Whaling Commission US National Aeronautics and Space Administration Results of Marine Biological Investigations Report of Observations or Samples Collected by Oceanographic Programmes Scientific Committee on Antarctic Research Scientific Committee on Problems of the Environment Scientific Committee on Oceanic Research Southern Ocean Co-ordination Group United Nations Development Programme United Nations Environment Programme United Nations Educational, Scientific and Cultural Organization World Meteorological Organization OS r- ewe M -H n u •-{ nj «0 <4-l E ■P O C O 0) C-H i o 4J 5 -H u O -P n v^ n] (0 ■H -rH 4.) > -P c C 0 <5 ffl Q> C I M X O I D -P -H I ■U (1) 0) c a VI Pi O 0) c e o ■P +J H C nJ M > D^ 0) -P I >i I S «J C * la o n o ■H 0) +J o P m 01 >e(ll(U(Xj-H-HOQ.' CJ SXT) (D-P Ul dtCO MTJ-P MOMWC C 0) E-f U 0) -H -H . to flj tji-H (0 P 13 [fl V4 C U JJ 0) X 0 -H < r^ c tP-O P >i- O M C J < C U dJ < w -H ; U a o c ^ -P J £ 0) , X c I &.H 01 -rt U O M -H ( « > <- W 0) O ,( >. 01 <-» M CO ^ ■H O > "W O a c o 01 -H 0 p tJl x: c J w c MO -■H >, O W ■ . . , QJ3C 4J4Joiua: 0) nj d) ct4 c TS COliOPtjI-iW c fO-ax: oJE-tstTifl u *4-i (0 a c ■H(0'aOQ)0) H -' oic n£rH4Jaj -p 0 p u-i J3 in . C^w -UOIt-tOJ-^'O 0 a> Q) s: o o x: ^ u w r- Qi4J ra 0 M oi u} en 01 V4 0) ' e -HO -H C4J*rH -HtJ^+J , C -P > -HO x:xiuoJcno'a'OH W3CJ-iQ.tOUaJi iH O 0) 3 V^ 0> A ^ -P 0) iH x: 0) ■r^ c x: 0) *-i - ■p (M > -H H ja -H 0 . m 0) u OJ £ 01 4J M « -H >i tr J3 c ■p -H M tJ > Q) •H 4J ^ 0 0) 0) 0) > IH > ■H t*-l M +J < 0) cS 3 (Oiu 01 ^ T3 0 3 rH 0 « 4) 0 U TS •H -W •W rH «»H « > <" B XI 0: B-O 0 01 0 V-H « K h U i 0 0) 0) 10 a B D g IS B U > 0 0-rt 0 fi « 01 0 +J -H V4 CU ^ 0 •u 00 a -H « rH «4-l >io 8 > a £ « B B 01 4j a V u a> 0 h ■H d 0) 0 C-H II) s > 4J IM -H -^ 3 iw .H lU > « — 4J 0 01 B b^ k40 ■o w c) 0 •<-( M Q 0 J= Z 3 *j 0. 01 H « 00 0 rH B H »-l 0 > U E B B-H > < B >,T3 K 0 0 « S -H •o 5 iH f-i O O 0 r^ -P IM -P 01 a « u 3 Q. « 0 U 0) (d ca » « 41 h— B i-H ■H •H -H • M Ul U •H 01 Ul 0 0 E ■P dij: 0 4J .^ <4-l iw H -H y fO a U 01 B S > lu O 0 0) U -P < ^ U-H J; « C 0 s h Q n) 13 01 £•§&§ — > 0) OTI ac B 14 n-H -H a 4J ■H <4^ B 0 •H -y 0) 0 ii a 3 - 10 rH 0 « 4J E on r^ 01 B 0 w -o +J ^ m 01 B u £ 8 >i-u -t a £ •H S 0 c 0. B s S 0 H-H ■H i x: 9 0 0) n ^i§^ M B JJ g M • >M a a « -H « h 01 E 3 « g B ^S2S s J= *J 0 p ii P n nj P < §. g8ii: s o a h •rl Q g 01 -p a V u XI o o 06 -H 0) E -H U 0 u 3 14 0 o *-• M-i 3 ^ 0 (4 a j= f-< B T) -H 00 V) -. Ul Di -O 0) tft a. c *J E 2 " ■•-< ^M t>A U) c c <-• >s h 3 4J -H M-( ^ •H *-' C > CO t-H rt v- a>-H )- c l-H 14^ 0 6 J T) 0 HH MH a « ■iH < C1.-M 0 Hi ■t-" TJ 0 CO-HX u 0 •H • CO 0 ■H *-! i/l OJ OJ rg 0 ■♦-» 1- *-» c e T3 4, rt C *j D CO rt C nj *-> 0'< c .^E s 0 ^ Q in *> r-l 0 X C 0 C t- CO C U 4> ^J £ uj CO £!.<-• 0 > rt ■t-* C :e: a C-H 0 CO 00 ■H Mh c 0 CO ^ e ti ■H e M )-( 0 c > a 4) C ■<-* dj <-» c s ^ 0 u > S »- 2 u V- -H Vh A) CO 0 ■H rt X i- ■ H V) jii 0 • <4-4 cx w CO > c C Jm ^-i 0) CO (D coo 0 S 0 Q S O tu c_j i 0 « J3 9 S J= 41 (» u M « (0 >, U C O J •H M CD O G 3 O OD •-H 4J « 4 ■H d p. < 0) (J 01 U M 4J u ■ O U >*-" o I -H O 01 (U I a u V n a a T3 ■H 0) C M-i -H 01 - O O *J ■»-) n xi Li -H (0 a -H a u o XI a, n CO to u 01 3 it-( O >. U 0) o at -H o O T-) a 3 4-* !-• i-i O ^^ 4* 01 Li ^ 60 -H OJ 00 C J3 3 -H TJ ■ O ■r^ « 2 Li U .H **-! * -ri -H o a • ^ iJ O !-< •H ^ « -rl « ^ 01 4J O 0) U L> <0 O U -H a M Ck •H 4J O O U U U •-( Li U U 1-1 p. a> M U -H « Ul B O 3 O 41 O *J M j= to 3 pq ii T) "H M-i 01 J= OP O 0) 4J Li ^ u-i -H 0) a 3 I t4 >H 0) O a ji o *j -a •H a 44 4J 4 TS 4 3 O U O 9 4 •rl £ U O OB a ID rH 3 4 TJ •H V 4 0) bO O « 00 >s O Li a OO-H G ^ a 1-1 U 4J 4 Cl 4J o a . a Li S i-< tA ti U ti 9i 4 t-1 n n > ^5 S-l >. a> O O rH j: G o ^ fl H -^ « XI o lis B !■ "ft S.E O E u.* O u s| ss <. d o 9 a U 0) tj ji 4 3 00 -H r^ > > ^ £ I a Li p 9 V Li 4 4J . ■Q 4 3 4 1-t 3 d 4 4 -H o ax o • -3 o U tA Ml 4 M 4 i 4 U Lj j3 -a u 4J a 0 U-i o 4 Li 4 -H O 1^ 4J CD S" ss H *<-■ iH 3 a 4J ^ O CD 3 Ll 4-> o -H o 4 a n E5 . 4 -H (J <4-l O £■0444 4 4 > a *J -O 4 iH i-H Li 4 U a 4 4 3 U O £4 4 1-1 O -r-i 4J ■H O 4 ^ 4 to ^ .O O > I u 4 r iH a ) «" o 4 4 U 4 4 4 5 ■a b Li 3 O. O 4 4 .^ O tXTl 00 t3 U 4 - M O > |ssg. O 4 Li O -H O TJ •HO 4 O 4 ^ U j: 4 q fX 4 4 ts 4 4 4 4 4 U « 4 4J « 5- J i-< > 4 t-i a o. 4 U j3 as 4 4 U 9 X j:> 4 O 4 u u fl >4-i 4-i>j::a3400 •H LI o 4 £ u 4 4 iH U LI a I S •> U iH < I j= j: -H L. U U £ iH 4 4 > W I 4 a Ei I ^ >4-l iH u q 4 •H O U 4 a 4 u a o 4 4 q o I a /-^ I C 4 Dd j:o4'HTjaoiH •^-'040 4 O i-l .C q Li 4 >>'^ ca CO LI -O 4 ■ V U M 4 4 4 V* H 4 4 U "H U iH 4 «J q u I iH u \ 4J 4 I CD LI g5 00 4 : fa u "•3 . 4 U « -H q J3 I i-« M q U O M 4 - U Q O -H 1-t d u 1-1 '-' 1-1 LI q U 4-1 .a 4 -^4 D. «-i LI -H 4-9 - >4 4 >■ -" - i IS O H 4 , _ _ I O. « LJ f 4 3 1H L j: 4 > o 00 44 U 00 -H 1-t £ 14 d 00 4 I 2 d 1-t o ■:» T^ I 3 i-( tJ r-f « o a 4 -ri M >s I - .H 4 iH ^ X M - O Li U I >~l 1-1 4 L> 4 d CO LI d O LI ■ O -H 3 O CL O f •H Ll 4 4 _ _ r _ , C ■_ 00 tiouKd oq 4-04iH "H"^ *L<4 .H ^ u 4 a q li O 1-1 LI O •H LI 4 U H ' *J -H O j^ ' 4 1-1 L^ 4 Ll 4 a O : 4 ^ .A 4 q -H o iJ a M rH 1-1 O -H q >sa o aLirHLJ O ^ S tL< 4 p> 4 a -H 4 4 M H 4 U -Q 00 V 3 T3 •»^ iH • a : jj H U tM V a 4 4 3 4 iH d >,£ U O a U O 4 fH U J= 4 4 N > Li 4 4 O. 4 q I' '-^ 4 d X 4 (A O L4 L-l 4 >M-* 4 M HiH^Lid3w.i. Q -H 4 O U iH O.H 4 4 > -H LJ 4 3 4)Li-abLlVd O 104444^40^3 H-aa4a4ad4 0) cr-H a) c -p c "H -H O 0 to M 0) y^ -p ui *J ■H fl] ^ .p to n a s x: 0 at sz^ o *J 4J XT' 0 0 iH O ITJ n c >, N 0 a-o 0 -H XI W >i 0) 1x4 a 0) 0 4J £ U 0 tP-P -4J 3 a ^ C flJ 0 0) • at 0 M (U e >i to > rH ia > 0) ■H ^1 -H u rH »W rH ■H rH iH •H -H ■H j-( .-1 at ■a S H *J a 0 w x: 0) O 3 £ 14-1 1^ 4J > +J 2 ■H j»; 0 ■iH to 4J >i rH c c D CL c d tP W (d » 0 a-H fl c £ ■H D -P 0 CO x: to H n tn to - ^ •H u 01 at tn c rH 0 J= to T) > ^ 0 ui (d m (0 rH JJ 0 'H D> -H flj -H at 0 Id E U u XI x: > ^w -C E 0 J< 0) 3 (d is ^ M x: i c E -H M >« cxjx: 0) 01 M Id S >iO tn c a) oj » H 4-t ja c •H XI -M U (d H ■S to ffl to ^^ • Tl u-i 4-> (d (d U en U OJ 0) 0 x: E e XI • u c M 3 4-)^ 3 13 •■ (U U) ro 0 to ^ j:: x: C «-< fN »4-< -H k< w 4-) O 0 0 H 0 u at 0) -t 3 "w ■p en w ^ •a 3 M (d 01 -H 01 at c ova x: i4 x: x: (d C D*^ Eh J<: H 4J , 0 T) 0) >i4-> 0) -O O to 4J M Id C -H to 0 E at x: M at -HO > M at 0 4J «*-) U t*-l rH J3 -H Id Id O -H IW J3 4-1 x: k4 c >4-i 4J M >,M Id OJ o 0) Di o 4J a t-i 01 u-i at u a. at 4J o a; to Id 3 J Id tn 3 u-i m v< CTi fd to 4J c o >i 0) to at tn -H cnx: at ^ to 01 ti t^ T3 Id 4J to c Ot ■--* j:: in a> 0 c > J at u i4 0) • o X3 o 4J tn M o ki 0] 1-1 u a o at Qj at o ox; < 4J 4J >, .. 4J xj 4-* (d Id o c ot T) 3 ^ in (d f^ j3 01 ■H rH k4 to O O^-H T3 a-H 4J 4J c -I C x: (d ■>-* u (d "H 4J O X: -P^ nH o 3 tn -o -H fo O OJ «*-! fH M OJ ' to > 0 oj j<: a O -4 Jj -H O ro id tn >, 01 -H c x: i4 x: OJ - O O 4-1 4J > OH *j ot a> ix> 4-> •• Id C »i-i 13 i 3 a> M M tp i4 73 cr E a 0 c ot c 01 e H x: (d oj cr 01 4J tn j: (d x: to to -H in H C 4-1 -H 0) M-i ^ (d > en E cn ot i^ at • -H • c ui Id x: 3 >i at H 01 jC 4J a) c ^ u 4J x: H ajrHutn-Pai>i> at Id 3 ot sz ^ 330>4-l4J^in 4J c to H (d (d -H oii::aj'dx:cox: aa (d M £ 4J -H -H 4J u XJ^ -H x: -H **-! ^ C 4J u 01 C 4J 0 >i^H y 0 T3 3 in tn 0 c u to 01 H 0 0 3 •p c tn fl V4 (» x: -1 S u at 0) x: to fl 5»°-g'8 fl 0) fl >i 01 T] <-« ^ 01 'P x: at *J 0) '-H 0 >, fl . 4Jumtnat4Jc otE flOI3-PC-Hl4-(l-|3 •44 TJ fl 3 0 P 4-1 0 c c 01 i-c 'H m 4-> at 4-1 V£> 4J 4J 4-1 01 in tn u .S!3„"S fl 3 0) I-I fl — x: Id to 01 iH at c (j^ oj 0 0 3 0) «> E-i E E « w 3 3 fl ■p ^ 0 (d x; m CL4J a43 .■85 4J 4J 01 • M to 01 013X1 'OflV40>, tTMO coi3aj-^tn CO ^ fl M cr4J >, >,u fl M 01 EO Id Cfltn«04Jcxa4-io 0 at c 4-» Ot a 01 ■HOj M-^tncEoo u a 0 '^ fl 0 4-1 0 a 0 0 'O>i0ifl4jo)id3i-ioi c x; OJ 0 > c fl fl«-44J>,3UfO -POI 4J0 -oflajxic^ tn *aiox:xaj5o M'or-tnk -H4-1 sz oto ocxat(4-i-<-(oi3 ■H M x: 0 3 C "tW 0 4J W 0) a> ^ c 4-1 0 ■H «4-( 01 fl H • at at x: ai -D x; E -H 0 U in 4-1 0 -H 4J 4J at rH 4-1 OJ 4-1 4-t OJ tn E 0 SZ^ at u ^ fl M 0) CO 4J -H i4 -H 0 >*-< 3 i-t T3 -P 0 0 0 3 4J u "w x: 0 OJ TI-HT3X:01X3 E-c C1-1C4JM 'n-rl4Jat 3 ot fl ^ $ ^ xi -4 fl 0 -H tn 0 0] H 3 CO a > X: fM M 4-1 >, tn 01 ■H 4-> >i- tn 0 -P 1 rH C c u M 13 c at 4-1 fl tn (X '- -^ tn tw 4-t l^ 0 -H H fl ot u 01 X: -H OH at -H 4-1 > -H 13 ■H ^O fl >|-H 0 "O 0 a to tri £ 0 4J 0 flT3 — -puac u ooiin -HiiOfl4Jat 0 id -^0 C fl u MO -H 4-1 fl 3 c fl to >1-| -P 0 Ot ■H 4J at +J > M -H 4-1 m at r^ >,x: ■H 0 0 HXJ fl at 3 fl tjv-H cnflatoaaiM-H M a---* — ^-H 4J at OE'O-^tT^C 4J-Hfl ■H -4 a.4J c OJ ■■-< a « a-H M to tn 0 x; 4J xj 4J 4-1 ■H *J c 0 c to ■p at u < 3 « TJ 1-1 tn at > x: 0= C01MX3H-H -H .-t 4J a; -rH rH ■H 4J >iO*M E a E 01 0 X3 '-' »i-( 01 X) 3 M 14-1 u 4J V4 M 0 at at fl at x:-H 0) OlTJXlTI -fl+JW 0 01 0 4J Ot fl tr E x: 01 J^ M 14-1 to 3 fl x: 4J 4J fl M -p 0 *j4Jo^4Jcat u flflC3 flrHMOd 3x:d04-i xiat4J> cr-PC33^id> at to • 0 0 M to E u E5 tn ■H c H c at H ^ fl fl 4J 14-1 at at x: M (H a E I 0 o» > x: P u at aj« c tT» x: at H ouu ■H>*4a.0tn fl4J4J'0 i3 4J 0) Id E-i 01 'H fl x: o-H 10 Id > c ^( ^ TJ >, 4J ^ fl 1*4 c 3 xt • 0+J i-H 4J -n 0 •H $ «M OJ 4J e >-( M U 14-1 fl C <-H at <-* x: ^ T3 0 0 > a-H x: fl at oia)x:aatotnoo3 >^ u u 0 H 3 -H a 4Jx: tp x:x:4-'a.3 h 00 ■H H 0 0 3 C ■P fl >i -P XI ■p4J-Hfltn>i ^ 0 tj> H 4J 3 w at 0 E M c ot 3 C 4J C M to at • x: 3 fl -H 3 m TS 0 fl -H 0 -H -H 4-1 to CT'^ fl S VD ■p Ti at tn fl oj OJ 0 x» to 4J H x: 0 C -O -H > -H U -H fl attHJ-l-H4->J^^-. 4J -0 C « 0 oj H 4J tn vn u at c > 4-> 3 c-H34-ifl •.oat fl 4J 4-> Oh tn 0 a-o 0 H atco4JooEai n 14-t 0 U 4-1 U OJ ^ 0) fl OJ to ■HO-HC 3nx:oi3 >-H4-> atTaTj o-p crfl at at to oio>'Px:3CE>-H oj a a ot -H 4J fl tn 0^ 0 -H O^C-HCO^w flO cototu-Hi-iccao £ X xx: x: fl a 01 ■H x: OJ 0 fl m E-i at 01 +J H 3 ■- M x: to J3 -p c fl M M E tflo-i a-H -H --' ja 00 =: 3 OJ >, O > tn rH I -H O -H C Jtf C ( W tH C 00 ■H m 0 ^4 = 0 fl -H fl 3 tn 4J 4-1 M T3 OJ 4J a: 0 tn in = o c C C XJ TJ O C tw -Hflatctn4Jflo fl o 01 fl TJ in o ■POM 01 MtOXrHOt7>tn4-> at « 0 3 c c fl o fl tn 'H o u E C ot 01 T3 -H «H o 0 tj^x: c 4J at 0 -H -H fl 4J fl fl x: X5 4J a 4J o +j tr fl TJ to o C 01 -P -'O •-i u -H Ij^-H tn fl 01 "H 0 4J M O -H MAT) tn fl rH rH T3 3 -H atrHafliHin4-'tn > X «+4 3 fl tn c Mflat4-toato>o fl -H x: e > o a: 4-1 «M a to m fl 0 O fl 4-» x: 01 c -p a to • 4J ot tn 0 3 3 JJ 0) -H -o E 0 TJ fl x: 4J at . rH0)M4JO0>.-^4-i to -P fl C W 3 fl OJ 'O c OJ j3 >i M x: -H ^ I rH4J4-10flatS « ■H m > M x: +J to 3 C "w fl 01 4-1 j^ i*H O -H X) 01 O c E 0 0) aj a> o •Hat>iEH*HE*^-p fl x) rH -rl x: to a 4«; 13 4J *j tn o . o> p^— c . cofl-Hcmt* oc [ox:30Qat'0 o fl cr-H a > 01 -o H Bco 4JO0lC0>-P O-PMOM otno ■H to fl 3 a" H fl 3 xioiajTJ oi-PXi-o i-i a o at 4-> c o OjO(XHx:flatoiM 3 "w fl a+j M B xa a o to to ot ■H M X) 4-t c • at to 0 a) D^ to ■H M fl H 0) •p « a U XI 0 -- 3 ai at x3 -O M to x; r^ 0 ot 4-1 4-1 3 M x: tn 0 a4J H x3 x: X c tn fl OM to 0 M * 4J 01 0 0 01 c 4J 4-1 0 3 H fl tn 4J ID 0 u m tn a D> -H at c tn >. x; H c rH 0) 4JXI -H fl J^ at -o M 3 0) 0) ■P C 0 M a) C -H C XI V4 0) (t) o -w s: o -I s •H 0 ■d > >, O rH 11 x: c 3 o o -u • C fl « J< J= ■u ei ^ OJ gg. £ I 4J U CO 3 C •O Tl H 01 O fl 11 M x: T3 a u c ■H ^iTJ u o T3 fl 0 c 6>H fl -H u u g a cj 0) £ ■H x; o *J o a> ^ o -p « 3 a> -p fl fl %t a o • fl >1 ■p • >1-H 0» ^ > -H C-rt li, o-p — "t " §■ I -H ^ a> a e at es V *-> OD V i i U > -H 4-1 B 4^ o M » c 01 0) 0) 01 00 u ^ 0) t-> ? j: M > M 10 o >-. iJ at 01 •o 0) m 0^ 0) B Q H) 3 ■H 3 n M U) 00 0 p j: c 3 o r •o l-l a 0) j: o (0 OJ o JZ a> *j 03 M 01 c o u < a. x: 01 Vi V x: c >4-l tj s nj 3 M o •o -• c 0 c u i-l c U 4J o 1^ 0} 0 to J "Q 01 •-) M U CO <: 1-1 (-1 o c !-• 01 B B T) (0 ■H ^ -H 0) ID T* <4-( O D O .-H O CN 01 W Vi i^-t O 0> CO ^Ei o a: o :x B « M U OJ U 1-1 41 CD U 01 d «-l -H ox X ' 0) 1-) M ^ a -H "u 01 3 iM 3 - I w d o; I k' M 3 I CD OJ O I O. «J c^ I 01 (0 I o CO d 0) M ■ a> X u ■ IX*-'*-' *j 3 w s O W 0 U-l to 3 0) CD u a SP o ^ o o M u a 1 3! td u *-* l-l X 0) 1-) OJ X l-* .. d a d o o O -H -H ■ O 4-1 (0 (0 ^ 1-. ^- l-l m d X kJdudAJWdcj o 3 CO i-i IJ 6D -H -H W O I- d a B M U *-> CD CO T^ t3 T3 *-» a. u < < CD < to OJ CO -HO* 1-1 a Q ^ M 1-i d w d 1 3 0) 0) -H Z 01 oc 3 3 a CO 00 T3 x: 01 o o -a o t-l < W W Ol, Dh < 4-t o Tj -U IT) -H C O Q) 01 0) rg «4-l C TJ tn 0 -P • c M tn u S > "w Q ■H (D U C • y-i a 0) 2 O O S 4J 1 c (4-1 rH ^H m ■p -p d C to 0 fd (0 > = O iTi U "P O 0 ■H 01 I 4-1 J-l J-l CD fl TJ 0 i-i QJ > C M rd C 0) C " Q OJ r- - a^ a x: TJ 1-1 Qj <-» r-1 0 rH QJ C E 4-1 = to CTi c E u E ■•-' (0 (1) -p ^ 01 M to fd -H a E 0 0 c fd C i w ^ ^ -H > M Q}Ti 0 ^ 0 -P ^< Tl 0) P to 0 U -P QJ -H 4J 1-1 m ra ip to .H . fO CJ Vj 0 4-1 -H u w QJ E 0) (w .P TJ tP p x: U (0 ■H -P 0 ■P c x: Q Q> M 3 C OJ RJ 3 C T3 P > TJ C ajj COS e c ceo TJ a C 0^■-^ XI in 0 -H 0 QJ C fd -H ■p 0 (4-1 l-l E O C Q) -rH H ^ CD ■H DC C TJ to P W --I W 4.) 4J 3 ja 0 E 0 S, -P M x: w 0 ■'^ O ■H TJ tn QJ -H ■-4 C TJ 3 C r-1 0 ■U 0) i c ■H -P C -P 4J -H -P 01 H > 01 > 0 0) TJ Q) M 3 C C c e Cm 0 -P 0 rH (U 3 0 fO -P 0) TJ W -H l4 c u to OJ x: to 0 0) --1 Q> u m 1^ J-l > c O T3 1-1 nJ OJ 3 Q OJ u -H x: 4J 0) Q) -P i cp - x: -H -H ■H (d C OJ 0 c (0 E E ^ U TI tP > u * P P -P e +J > rH 0 u >t at a !-< U OJ QJ c .-H c a> to ip to QJ O 0 K CU c o c a u c w .H +J OJ 0 +j c ji: i-i fd rd -H P 0 to in 3 fd u u V Q) tytA U (C +J 0) C 1-1 (P fd -H 4J C TI OJ OJ +J dj M Qj a) ■H -O -P c +j -H a c x: 0) CP ■H c OJ x: en u c 0 TI 0 -H 3 ^^ c c -P +J QJ W 0) fl -H +J 4J 4J 4J c «-i Q) x: 4J C 0 OJ x: 0) ■p ja 0 a T] n-t U V} ^i 3 -P c 0 fO C ■r-t D.E- ■P P E 3 10 •H W TS 0 ta ro 0 cro-t+iu-icj=, E a 3 O >i W 0) 0) -P -P t^ cu a tn 0 OJ c c 0 -P E OJ T) iP ■H O OJ Cu -p in cc; O fO Q 0 u ID m d to E QJ ip Q) • -H 4-1 C 0 to 0 ■H 0 0 x: >i - U 4-1 (0 TJ -P -P QJ C to O 0 >itn -P O TJ 0 u c cx fd 4.) a o s ^ fO 0 4J ■H XI n] 01 13 0 XI 3 OJ 01 >i^ T) QJ x: cx^ -p a U H-t J-l ' 0 E -P -M OJ T) U 0 c to XI W -H 0 ^ 4-1 C 0 tP C 0) 0) CP 0 0 ffl (0 4J w ro x: c o -p td to -HP P 0) OJ 0 ■y c 01 O l-l C U-i ■H -H >i 1-1 x: ■P fO M u M Tl > td x: p 0 0) C -H B C CU ■H C E -H 0 d x: a u IJi UJ c U QJ :2 U r-^ X: rH c +J E -H owe ■■H C tJi M +J OJ nJ T) c -p 0 c x: . 0 u -P C -H CP QJ (U ^ U (U +J in nj OJ JJ Q a C ■H U -H OJ ■P +j x: 0) ■H fd C £ ^ M -P TJ -P (0 -P c w Hi e tl M rd en M c to x: -o m u 0 nJ U ^ O -P 0) I-l 3 Vj Q) -H 3 Q. tn QJ fd W Q) 4-> 0 0 y-i -M -H d T) -H j:: '-^ 0 >.< ox: Di T) e -H iM O E -P 3 +J TJ 01 u 3 Ui +J M C L^ -p w a t-i y-l 4J ^ C ■H E OJ O C C 0 .-t C 3 0) Q) 3 U QJ QJ 3 H E rt C OJ fO -H T) E S-i to ^ QJ 0 x: f-i di >-{ E TI 0 r-H l-t 1= € 01 U -iH C -H XI -P C -P 0) ^ 0 ■p a E w 0) E O c >n (0 fO (U E O 3 Q -H fO 'H O a (d u j^ -H a 0) in 3 Q) C fd fd P ■P -P tr 0 XI D— 0) E 0) ID x: VI o 0 -P -P 3 0 4J 4J tn 4J -P p y-i M C C u -1 E ■H tj C -P E -H C OJ H to to td c M to fd U 3 C O < fO OJ -P > -p dj .H IT] (D T) QJ 0) x: 3 01 4-1 0) u fd 4J TJ OJ 0) u r] E )^ c C 1-1 0 ^ tJ- C (0 > E -P p P 10 E Q Ul fP XI -P •p x: c 4-1 m 0) - (U JJ 1h CL OJ C to 1-) 0) c c o E to 3 c > 4J E-t 0 0 < cr £ 0 > fO CJi (TJ (d -rH fd ax) c +J Q. OJ c fd C -H IT] -P (U jj jz: (d t^ to (-1 (d QJ C Jh ^4 10 PECO) u E -p x: fd Q) U -P x: in v^ to -P 4J nJ: £; a -a u 4-1 "5 1:3 OZ Ed S X > ip Q C O S W fd -M c ■P y-i C o o fH O 0) CM +J .H e CO bO •H QJ V r- C •H nJ E -C ■ ( o to fd -H o o f4 +j c x: TJ fd'H p^ +J > o Q) t. TJ C to X; 0) QJ -H fj +J to 4-1 E 01 C W T) Tl O OJ O -H 0) U 3 3 to 3 cr+J c 0) Sh OJ o ■H O tn f^ O E4 QJ ffl to '3 " ft (d bo (0 OJ x; 0) 01 o PS O X o td 3 •H O f4 P4 W • OJ O OJ CD 4-1 a-, a; t^ en M 4-' OJ f x: E t4 -1 4-1 ol fd tp tl S4 O -N bO fd C -H > XI QJ fd -H tU E +J -J U- •p fd 4-" QJ > 4-" -M x: c c OJ V O o to P to bO W CJ -P O tj fd o w 4-. >,-p ■p C XI XI ■T3 < x) to •• QJ QJ +-> ro x: -P -P n 4-) fd O TJ TI x: o c C bO-H fd fd 3 O QJ (0 fd fd QJ TJ u c fd 3 O Q) XI x: ■M OJ x: bO+J 1 +-" -p CO x; xi 4-" fd u 4-' C ■3 01 10 nH C 01 cr a QJ < td 6 0) X bO o 01 TJ 0) fd U ■d a • 4-' 3 TJ QJ -P to ■ +J C C 6 fd c fd O -P " QJ 0) bOW E E C ^ (U 0) 3 0 (0 t- -P o a -p C Ih ■•H 0 -P o 01 fd x: ■ Tl o ■p x: > a. ■p 4J 4-" fd tU TJ a c u ■ x: yj O >^ 0 bO VH XI -P o t4 4-j -p c ; " 3 c X) -P fd 3-I-, QJ 0) (0 OJ 0 E S^ O O B 01 a o ^ f^ Q) 3 bO > XI O fd O 01 rH ■ c o o fd T) C fd c t:i dJ-P S 01 OJ -P ■ *-> x: XI bo to 10 4-' 0) u >^ •H -H x: o QJ x: -H 3 H i^ > a. c t- fd OJ 01 c E C I >^'p I U 01 fd c ■p< QJ r-l 3 x; +j OJ to 4-" b04-' C 0 fd C 0) ■P CX td E a Q) bO C-P 4-1 c o o fd •p -p P" C Td 4-' to -P 0) t4 td JJ td >H +J to D. fd u---t a QJ rH SL, -P a, -H tp 0) 0) (0 t4 x; 01 4-" fd 4J x: C P QJ (J TI E-P C ip QJ 4-" fd O 0) o i^ U *-> io bO td C tn (d 4-» QJ QJ C E 3 P < 5 QJ X: QJ -H bO QJ ^ > ■P x: bO QJ Ul 4-> fd S4 1 -P "O X) >i x: ■H T( 1 0) 0) c fd 0) -H >W l-( (d u 4J QJ (d XI 13 C ■p c x: 0 -H TJ > - 0 0) ■H T3 c fd N 1 E 4-1 M ■H -H (d -H 0 -P iJ u c 0 to x: to x: n-l o c ■H 1-1 0 4J fd 01 TJ > -o >i x: 3 -H 0 -H 0 (d ij QJ c tn fd 3 M 3 ■H J-l 4J C QJ 1-1 fd T3 -H 0 -H QJ 13 C 4-)ii-)4-t-Ptj' >waau o-H 0 0 4J QJ c fd t: 4-1 j: in --4 3 1-1 tn > ^ fd 0 OJ o [?> fd c ty c w X t^ ,H -H iw X) "P M-l x: O 4-1 01 CP C 3 ■H tn 0 QJ a a 0 x: J3 > ■>H CC-Pdt-H^ (UfdOJ 0 3 fd -P Q) 0 to 0 -t 1-. M 3 10 c x: > ■H 1-1 0 -P CT' -H -P 0 -H H *J C XI 4J 0 -P O - JJ c in fd 1-1 0 l-i T) QJ 0 in -H 0 j:: ex, -P -H c x: -p u ■HC0fdl-((d01J'0(D W C 0) 3 o tn 0 c 3 C QJ C Vj U 4J +J tn S ■-H U (d fd ■pfd^^oi-HjJwcxj'Ocuai'Hoa QJ -H 0 tr u >i fd 4-1 (d at c fd c w w TJ fl rH aoiatro-H i-i(d QJ M E -H M T3 > -P C ■H C C 01 -H 'U x: in o c: TI 0 Q) a4--H D OJ QJ 13 EEXOCIUMQJ T3 C -H 0 QJ C ■H fd 0 4-1 -H fd U U Cr^ - 4-1 3 -H l-i 0) -H -H -H (d .H M OJ 3 (u'aoiHO>W'H {Ji Q) I-l U XI 0 ■P E •r4 u TJ 0 -H T) C 0 10 QJ -o -P TJ j:: oj O -P tn tfjnj fdu -H'0^Ti QJ O u tj Id a CP4-' tn M l4 3 ^ fO ^ ja c; --t x: j:: aid-Pfd-Ptoidcu -px: Q) X 1^ QJ X) ■r-.U-i < T3 at to -HO at OJ u fd Q. (U 0 -P ■p -P 0 C'HOCx: -uccn T3M-«Qtotncjac 0) 1-1 c 4-1 tn 0 x: > in in X M )^ H (0 (U u at QJ-P C 0-PT3 -H-H 0 4-1 tn fd O ■'-t xl to Ch 0 U H 4J 3 U o -H to oj 0 -p x: U W T3 u ax: e c -h o o id oj QJ " (d 3 QJ 0 ■H QJ 0 at tn TJ -H nj 4J ■H C C c 4Ja)fdtnjJ'aN>jj)-i E C T3 Tl O T3 U "4-1 to a at 4-1 r~i c u s: c M -p s: 01 in >irH 4-» 0 fd OJ 0) rHAJ-HUC:>l(dM(U -H 0 QJ C tn 0) 0 0 0 o x: o 4-1 0 QJ 01 0 -H QJ 4J ja 0} Ti oj 01 U-H D x: inQjOl -HfdrH QJJi: CP-H ,-1 (d -H c cu u 4-t QJ OJ-rHTJ ^QJEl^x: ■H 0 M OJ Ul 4-) - ■p ■HEJaCT! (d >i 0 a in 1-1 in fd Ti ja 01 H 4-1 0 -P J3 -H Id 3 -P iM M fd O -H E T) 1-1 c to Ot QJ . ^ ° tn -H ^1 0) - in 0 fl^ 01 ■H in 4-) 3 M in tn rH ,-< -P O 0 C -H fd ■H 0 tn 0 -p Cu---\ u •-* fd M -P x: x: Qt at ja x: ■H ■H U 0 fd 4-1 x: • c (0 'H ffl ja < s-< !-< uj-PUfd ■PT3a)(iti 4-1 4-) » JJ T3 -P 1-1 4J 10 X: rH 4J rH CO >i fd J= 0 c o -H ^ 4J 0 XI -H 0 c 0 -P l-t rH 4-1 (d 4J in c c a-H OJ 3 OJ fO 3 C X5 -P C 4J U H H tn fd fd cn*M (d 0 xJ I+-1 fd >i u >,-H a^ O 3 0 c 3 -H ■H OJ 1-1 1-1 Si u 0 0 C 0 -P w QtSCC-H^iDCO +j H 3 a fd Cf 4J at u 0 x: o tn >i C QJ s: d d 0) 'H <4-i Ot 13 ■-H 0 3 (Ji-H -rH M a. C -rH QJ ■•-i U Q) fd -rH Q) 0) in Ti a4-'4-l-HX:rH OTITJ WW x: -p 0 > -H r-t u O 3 Q> X -H -P l^ r^ 0 > C -P C TJ to ■H QJ 3 M -H 01 -H (U >i-P n3 ■H 3 O in -P'DO-PM (UfdCXtOt ■ H in H 0 W -H o (d QJ 0 TJ HD 1-1 1 4J 4-t ■H 4-1 in tn rM 1-1 Tt C m +j 0 a ■H to fd ,H C -P 0 Cnx: XI JJ 'H -H c > o 3 O 1 -H (d ' 3 a tn c 0 TJ M Qj fd 3 T3 01 i c u ■H in o 3 01 in o u cn 0 (0 -p i^ 0) >i 0 QJ fd -H 1-1 u-l -H (d 0 •rH -O ^ C 01 c tn (d S-l fd c E M H > 4J .H in fO >i c ;3 1-1 o u x: CJ OJP +J01-PC-H to OJ ^ 3 l-( XI QJ a c c fd ■H W "4-1 >i-H U (d -H OJ a c M '-H 0 x: U-l 4J l^OtEuincxi-'HMcn at 0) QJ -n 01 E 0 0 M c -a (d Q 4-1 -H 4J fd C TJ TJ 0 fa 0 in Qj JJ en cn-H -Hid 0 [0T3C CT3T3 ■rH r-\ H 01 ■r4 0) at -H 4-1 (d a 0 3 c -H c ox: Q> x: x: c T3l3i+Jm-P-P.H 3 tP 0 C -H U QJ tn 4J E ia ja tn Q) T3 U C E -H OJ rH c 0 JJ O -r^ 4-1 1-1 -P s u -H inocufdfdtfltna • o at c C U D 3 QJ > H .H ■H ■H ■H ji: fd V4 ■rH 4J XI CJ 0 ■H ,H > 'H 0 fd 4J.Hi-ix:xixi3Xintnx:o-H o a QJ U < 1-1 4-1 4J >i X QJ ■H C H 4-1 fd 4-1 fd -H 0 (d en X x; U QJ > C OJ 0 ■H a,ii-HH4Ji4-< 0 fd GjOJ QJ E e Cn4J C T3 4-1 4J TJ T3 4J -H O C 3 +J 01 Qj 4J -H x: -H 0 j«; .-H fd -P o-H-HEx;'aM^^C(dcD QJ C fd 3 w ^ 0) at 4H 1-1 IM W ^ --H Id Q) ax: uifd-Ptn ^jjj.Hjj ■H 0 QJ 'H X) E ,H OJ OJ >i 3 x: -H en fd E C TJ TJ 0 M C -H -H tn E U E MC>UfdtnfOc in M-i a4J a. -< XJ E tn -1 0 en E C --^ C 0 G QJ (u rd a> -P -H Qj in c M Qj-Hi-ifd -PEfd-H w C C to T3 0 0 01 < ^ x: 0 0) 4J -H 4-) 0 fd TJ OJ x: Qj 0) c XI C 3 -P (d a,-P(UM-ja)OM-Pid(n 0 -H QJ QJ Qt rH M Xl CP 3 to 0 -H -H TJ C IH P C in ■P >-l 3 0 ra ■H o x: +J into >(dOtn4-»^ U ,H > 13 QJ 4-) -H fd IM x: XI CO) -H 10 c fd u U +J -H *J 1-1 t/j tji m x) -H c QJ 1-1 ay-< XI c in at H J-. QJ > C 3 c TI 4-1 0 TJ Id 4J > fd 0 a 3 1+^ at u w (d -H -H H CXOX:Q)EC3Q)h 0 x: x: Id QJ at 0 0 fd QJ C 'H 1-1 C 4J 0 C QJ -H X 0 o fd ja w cu E -o e -u fdQJUE-i iH tn 1 4-1 tn OJ 4H 0 13 0 H G at fd OJ 1 1-1 0 E d 1 4J QJ -H Id fd 0 M rH G tn 4-1 )_l 4J UH fd fd Id 1 fll .H 4-( ■H H C r-t •'-i CLS: r-^ QJ H in 0 3 0 QJ H in fd fd •- 0 > >< 4J OJ _n 4J to o cr u 0 fd UO4JT10J3 Mfd 4-1 QJ 0 -rl -H in a x: i4Ti M>,Mxi c tn 4-1 U E QJ fd otniH -hm aix:aEj:^ 0 10 QJ x: 4-< 0 >i H H>,4Jl-iO0J4-lQJQJO 0 ,—\ fd i-i 4J • IH E OJ ajM4Ha>id4-io34J0j fd 0 0 IH tn 3 QJ rH 3 JZ ^ 0\-r4Gtnx: 13 u c QJ QJ 0) U Tl TJrHGO-H 4HH aE=TJ u a4J 01 4H XI a4J erin4-iin T)rHQjd4Jdaj 0 XI TJ 4-1 fd 01 OJ u daat4-iuoj-H4-(i-i -h 3 a 1-1 x: in 0 -rH 10 0 0 GGQJ4-idaEO 04-1 C -H TJ at a to x: 'H 3-HCn 0Jx;4-iQ)QJQJ4-)l-irH 3 r^ c ^4 at cd ooj-riwfdaaJOxr-HO 0 4-1 c 0 in rH E tn 4J 4-1 00 -a-PdTJx:'-HaooT3 a TJ 0 d 4J x: n ■Htnoat fdo 04Jat ■H fd fd 4-1 0 fd -H OJ u indO'4-itn QJ 4-ixjo d G Tl at -H 0 in 0 -H 4J0JQj>tn i-iO)-Htd4-i 4-1 3 a M tn 4h IH -HOrH aJTl4-lQJfd =■■-( Id 01 d a 4J -H 'H fd TJ tn (diHasHQjQjox:xrH4H u •-* QJ in 0 at rH tn 0fdx:M4JQjx:xi-'H0x:d -t3 X fd 0 U 4-1 TJ QJ d OJ •-i aio fd-HXi4H4-i3 3 fd fd fd u M 1-1 c fd fd 4J 4-ia too GS-HinOJO in 4-1 rH fd 'H -H OJ x: 0 c a > 0) (d a QJ 4J QJ d H QJ4J(dOO (drHl-l4-) QJ QJ in -4-1 4-1 Mx:'4-j ajTiQJOJ«-io>i rH (U X QJ CT- c at to < 3aiV4-H4-i4J4-iintnPQjat > >tH >, 0 IH -H 0 0 >i arH 1-1 0 arH c fd a QJ at XI 0 j:: fd 4-1 -H -H tn > tpTJ a 0 in 01 1-1 3 V4 0 > in-H jain30)3 4-1 0 iH rH ax: ' E a4H 014-1 >,fdTJin 3QJdOJ H QJ xJ Id CT 0 10 ■H atx:oj o>inEx:o H at 3 fd 4-1 TJ d c u o O fNXI OJrHs CtnrHidTJ u d CP to 0 4-1 4J >i 4-1 -H3Ul34HX:-HG01OQJ • 4-1 TJ 4-J c fd 0 fd 3 C - CV4rH4JO T3d G 0 "i >, 10 fd to -H rH 0 u dOJOtn4-iaJ4-ifdiHio u ot tJ* O W 4H O l-« O aCT>fO= TJ "tdQJtdU-HE'UCfd ■H -H a rH 01 -H x» 4-1 fd 0JC0fd4-l 0 lOQJ-HQJ <3 fcn C H M 0 iH H dTl .C= OTJ4-i0)4-13rH0JaiH • +j rH 0 x: fd 0 aMTJOdtnajo>i ti-h 'H to w ja > tn 01 -H iH.fdG -H 4H10EQJ=X aTJ a d 3 at " x: QJ TJ tnodQJOQJ4H4-itn4-iC4-i rH 'O c c Q in G QJ x: 4-1 0- OQJint04HQJ-H-H QJ 01 ■H 0 4-1 G QJ rH U QJ 4-1 3 44 -H Jh 44 0 0 -rH -H (d OJ fd fd rH C (IJ i. 4-1 in OtnrH-HGdMtd34J>,0J c in V4 r-l fd 01-H *j CnOX14H+J3QJTJO > M tn QJ a QJ 'H -H -H = QJ oat 4-ioow era >qj 0 3 0 x: 4J QJ a 1^ G TJ fd dfdfdfdfdin OJOjini-i-H at 0 E X x: fd QJ 4J > fdotnid OQ>i oojoja -H in -P 0 x: E 3 -H ■H ■H4H OfdTJC 304J Tl a OJ 4-1 4-1 4-1 x: fd 4-1 iH -H014JX; ^ ^ •-* Xi 4J in 01 u C 4-1 fd 4-1 TJ ot 0 c T) >.-r4 QJ d ep QJ 4-1 0 01 0 1-1 QJ a-p c fd fd C -P CP-rt oinoJQJid4JXlfdi3 ITJ E Tl 0 X fd 01 ^ 0 i-irH drHMH E fd-Hx: fd >ifd Em U 0 rH X: TJ QJ 1-1 s: -C -HUfdOH-HXiniHfdfO ^ > u 4H at 4H 01 d 01 tn QJ(dfd4->-H [n4-l4-lrH a4H Id 4-1 U OJ QJ 4-1 fd arH X 4J )-i 4J c 1-1 3 U 01 0) MO 4H TJ in UU 04-IG--QJ in4JTJ T3 ■H ■H QJ X iH 4-1 >iajVH as-P otdin-HOO QJ 4J x: 4J fd 1-1 QJ rH fd c-HdQicodTJ'O oat OJ QJ 3 4-1 3 X: QJ 0 rH TJ d oacx (dTi>aicfd x: in 4J OJ C 0 X: T3 3 otnoi-iQj-HO daiQJ4-i to J2 0 G 4-1 4H fd QJ fd fd oajox:cTJoo4-i *« G HH in in 0 4H 4-1 d 0 u 0 >tH -h 13 4J -h . fd x: iH fd 0 4-J <-i fd E * QJ 4-1 rHTi i-i4-i3tdid -Htntn 0 0 JG QJ -H fd 0 0 X: 4-1 T) -H (d 4-1 0 4-1 -H -H a <-{ 4-1 0) 4J TJ X3 TJ Id 4-1 adx:aj =fd4-i3rHG 0 4J to tn 10 - d da3d >(d4Jin TJO 0 4h fd to rH Id aj G 4-1 fd fd4J>£C' fdinojo in ■H d 0 w Qt 0 - -P epo ii-H.-)H4-'ajccp u 1-1 0 •§ XI x: Q-TJ fd tn x; OJ lHOV40C'-rH >H 4H G 3 0 a 3 T) >i fd d ■HMH inoic oi^iojin Ch n 0 4-1 0 QJ 4Jjdtno= QJ o>i>.3eojtn 0 0 ■H M 0 3 d 4-1 x: ■H 4-ioiHiHatinQj - H ■■-< Xi Vi C -P tn U rH d G in fdaj4H 4-itn-HrHrHa3rHtn •H QJ 4-> 0 (n r-i 0 -r^ iJ 4-1 fd\4-io-HCEio-PiH fd 0 a 4J OJ -H 0 ■H 01 4JrH Tl4-llHtnOE 3 CP+J 10 0 a H OH > tn ETiin 4JO0idfTJOi3 ■H rH 4H 0 > 4-1 -H iH a4-i ■fddfdfd3a-HTiu C -H fd fd TJ C -P H G at J-. C -H >,-H U rH 0 rH 4-1 rH H tn -H 0 U fd 0) QJ 4-1 4J 3 tn-HQJrH QJ4JrH0 XrHtn -H G 0 Qt -H fd 4-1 0 > 0 fd TJ -H > a-H 3 -H 3 0 tn fd -H 4-t TJ 1-c U -H tn QJ O to 3 4J a-r| -H -H a fd 01 -H 4J -H OJ tn QJ E 0 -H u 4H 4J-Hx:e-pado\ 3 4-1 Tl tn fd G Od 4-iOQJOE>(Ue-H'0 tn 4H OJ x: QJ X: 0 ^4 3 4-1 fd drHat04JO-Hfdoo-*ji3 0 OJ at fd QJ C OJ x: 01 M-HtnendTJrH-Hjaot >i ■H Q) j::4Jx:e-i4JOti "is: ■H(d£Qju-H rHaE G in TJ QJ XI x: QJ Xa T3 iji 3l4Q)G CUlfi O X - QJrHTJ 4-1 4J 4h 0 rH U4JV4fdj::-3 +J(0 ■H c 4-1 QJ QJ 3 0 oao-Hincpx tnojx: T3 G IH 3 >i QJ-H -H 34-lCnrHT3fd TJ Ot ay x> 4J tn 0 -P tn -Hd-HdO) OrHrHH rH to tn QJ x: . QJ -H a axi rHCPCPTiTi CQJrHCXiCP ,-{ c £ 4H ~ tn b Si 0)d4Jtd -rH ..4J0jja 3 fd ■H E tn at TJ 0 XlOG QJOQJMfOfd+JG QJ 4-1 ■H -p o 4-1 3 a 4-1 01 J-iOOQJ -d^-iTlOJ >fd 4H -H ■H M G at - aTJ Id rH ^H QJ 4-1 -P E -H X 4J 0 e 0 i-i J3 ■HfdEotfdOiHUtnatd • tJ^rH to CP rH 0) fd rH >. OJ ■H04Jj3id a-4H.wtn4J E-< -H in - 4-> d iH < CP4JM rHQJ fdOJ -H -H-H in d rH < QJ Qas: CL.£i 4-1 iH 4J rH-Hfd ■Htnoto04->QJtn rH ■H at rH x: a 13 cfdaojaErHTJx: fdrH ■H QJ M E 3 X fd -H 0 0 QJXlrHTJOAirHid M-HQJ iH 3 01 U <-i .H> xiE atd3rHd4J0J >tJTi fdtn>fd>ifdOfi> d 3 0 QJ QJ H to \ at U 3rHOOQJ4JdfdO> 0 TJ a) tn t. -H QJ 3 fd QJ 0 tn rH C 13 MH QJt7i3tnO>OOaOJlH n C x: 0) (d x: x: 0 -HQJdOX4Ja)4JGO-H301 0) to x: 0 rH QJ QJ 0) G 4-1 4-ij::oJoin4-jQj3-H afd 4-1 fd 3 ux: 3 4J 3 -H in fd4-'ajMrHtnfd 0^4-1 tnrH E ITJ 4J fd QJ XI M TJ lU fd 04-ii-iUfdtni3er4-Jfdtnx: CM 0 u tn 'O OJ O 01 -H Q) " Q> 4J 01 M 4J OJ QJ T) H M 4J x: "4-1 x: c .^ Q)TJM 3 *nj4J^ 4-1 -H TJ cn 4-1 > x; C0)cn4Jcn cnQ)4J'OCL o) 3 fd 01 QJ >i QJ OJ a-H 3 l4 01 4-t OJ Q) W o»c3Cfd ■r4x:-H^-Hy-*{n MOM tn XJ QJ M "HOC 14 rH x;huqjo'0'Hx:4J 3uo4J Qi 0) -Ji -P in- X30 'OXl4->o QJ U fd M e QJ 0 OJ 4-'tam-HEC 4-1 4-1 4J vj CC-O-HC 0^0>Wl-iO - OJ 3 x: 0 QJcnoc4Jcu oco ■H QJ cn IT) [/} = 01 OJ >i ■ 0)0 u-io-o-Hooi-H (a-r4ai -H 0 4-> U w-hec Oh "Mnj 0 4J TJ M 4-) ^ C OJ fd (0 0 x: >i x: x: fo 3 U(U0J«-I-HCU4J ^4J4JQ44JE 3 cn x: )-i'Di4 (d in-H cj^cn ii>m Q.4J in 4-> 4-) ~ " 4-t t O 4J x:34-'Ln-r4x:y-(UM iTJ-H 0 OJ 0 J-< u 3 OlUOtMO'd QJCn 0 0 fO Ul .-1 cnw4J[nn3(n'OCLQ)OJ --t > Jj^ C M 4J (d 3 OCOX: 3-H.HOIC.HOt 4-1 QJ U ^ i) x: Q) D " 01 c c>3'a'd)-<^M ^i-(0) M ^ cn oc4JC'H>oi4(da)o (d > OJ 01 4-1 -H <— 1 M CL,f-i OJ 4j -o tu crxi 0JO4->-Hl-iUff]l4 U-t .POJI-I u tj" q -H OMOnJO-HrdUt-iO rH ■H " c Xl-H(o Qjcn CPOOJCny-icn fd c 0 E U-t -H 3 -H > M XI 3 cn 4-) Xi 01 13 -H (U "X) > ■-< 13 4Jx:cntn-H0j(d4J c coi -H -H -H ■ 04-J[n4-t CL CT'C/imW 'H V4 x: 0 ■ ^ Q) 'aa4-)-HCT)x:u( aj-H4Jox: TJ > 4-* W « CL U OJ ■-( C M C 4-J OJ C 3 ^: 4-> 0) !-l 0) fO U] ■HO 0 4J(d'ax34-'(dcj4J r-4 -H tj OJ -n ii)_c:tntrt-^[i]nii_i m n .-J 9 , . CP a P Qj in 3 TJouo) acu cnx: 3.H0JX:U^ •4-i4J0J-H'DQ-Hbr'c^'n . ;$ i-i 1-1 (d - - - W ti 3 oooiui ci4J (i U x:34Jatooo4-ii-i4J> s: T) QJ 3 4J r-i a M E avi x: 14 0 XI 3 m 4J QJ 0 MU-(0'-104-l -. T^x: W 4-» 0) -iH c u x:qj4Joo-h u '4-IC4J C ^ OJ M QJ oi4 ffj M(dy-i mx:h-h 0) OJ 3 i4 cn 4-1 1— 1 0 cn 0J4JXlfd0l 4J0J OJC -rA c OJ -H 0 -H O CL4-i-r4_cn(nmfn— J E > C U QJ fd (d M OJ 01 c c wee > xraojfdxiajE-Hmj-icu >4J a-CT'M>i OT14J dJBcoi 4J > 0 OJ TJ ■H 0 ■H4-<'a4JWX:i-lHXl-H 334-' -H cn OJ 3 c 3CM4-) Ml4004«; 4J X^ -H Ot V4 e ^ x: -H 4J O -H 4J CD CT"a 0 O X 4-t 0) 01 0 H Ofl-H >iOJ'd(dx;cj-H (d w TJ 0) 3 = CP4J fd 3cniU TJ 1-iQJiD-HOlOJ rd > U U > cn x:4-)^Eoj 4J .H 4-1 TJ -H (0 j:: o • 1;^ 9 • D Q. CC04-IM.-HH •OIMCT'>cn C J-i cn -H 0)1^4-" fd-H-H l4 CP-H tn M "H QJ U 0) C} fO 3 • O 0 V40x:-H(d^cn34J (dxa oj 1-1 (d (y M ^ 'i4 0J x;3tj'idC'd3at 3 4J E TJ 0 H 0 fU 6 u-i C 4-) • x: Qj-Hcnexi(dco4Jcoiocx: OJ x; T) -H 1 'aw4Ji4-iaj -H Ox: ■^ ^ "tl 9 cn cn >i4J i-l 4-110 Xi O-HOOC ■H4J ■P c tJiCM l4CT''0''-H4J C3QJ0JT3 C0J>il4 rH 3 cn E 0) -H c 4-' -H Oi 'Hu]4-)n3oouojja-Hajoi ^ ^ E ■»-' 0 0) -H 4-) to X e rtj 0) 0 x: -H 0) (0 x: U-t 4J 4J < -H 3 -H C fd E ■H i43'Hc-H0J>-ia3'djacn(dx: >cn'ai4304-tU(.-(>C cr!iJ 0 d 14 -H x: E 0) D 4-1 4-1 (D - 0 **-tcn» x:cni-i'ax:>tu -h *M -H cn 144 -H0t'a4JO-HOXl'd "3 Cu vi t:: O-HCOJOOJO (l)CPl-l4J4J4-)>, 0 c X cn 0 •Htjid ,C4Joi cot x:'4J 3 4J cn TJ 4J +J CU - O O O tJ 0 IJi'H ---tCLWHOtOOU— 1 0 (d QJ 1 i4 >icntd4JcnHx:>i 0) 1 0 x: x: cn a OJ 1 <4-l -H 4-> c ■Hfdx:'HtitnEcn4-'ajQJC 0'a4->ai3Q-33 ccn-Hii-io C E -H c 0 C CO C3'O.H >On3&H4JCT' OOWXacOUM OJ QJ C TJ " > fd QJ 4-" 4-1 C a. Qj -HOJ-H Q,tnUaDO y-iy-i -H 0 (U 3 H c cn 3 Id M 0) cuxi u-i 4J C l4 4-1 i-l 0 4-) X3 cn C 0 ^ tn.-H'OCC'd cn o •-* o d) tn cn -H x: 4-t ojoxsbjcn u • 0 1^ 0 CT'--I C fd H 1 XI Cn-rHXlOO O-HOJ rHUi cu CO 4J 4-1 E Id '4-ii^x:oQC'a3cn u H X^ > 0 iM C QJ 0) cn^ (d 1-. 3 U 3'dtd Q-4J4-''OOi'd(0 cnu O -ri 0) 4J 0 Mi4 0Ccnx;iH4J 4-1 -H Qj -HOC TJ tn U4J C3C (d 0-HM. U XI M 4J -H cj> cuoJUid 4-11—10) Qti-i 4-1 Qj 4-" C C 0 OJ Q) (d cn 0) CO OJ C 0 Q) u in iit*4 OJ tn 3 oi M -H QJ cn 0 c c CO x: 01 fd fd 0 fd (d 3 OJ H CL u) e at e ^4 > 0 4-" XI E u-j ■H'OMHCE'dOJ ■HTJ>Ha)0> ■H x: u-i >,^ 0-HMcn4JfamuQ. m -H c cn 0 4-» T) Q) TD 4JOQJ0J>C 3fdEcn-H -o 0 ^ cn a-H > tj H c c ^ c l4 (J l4 c >« c n 0) &.-H 4-1 Q,H O 4J iH ■<-! 0) 4-) 14 O 0 X 4-1 -H Q - >i -H fd >t (d -H Oi u ni fi) 4-> 0 fl 0 Ul 4-1 x: 4-1 cn 0 1-1 0 o< fd o OJJ-i •04J(da4-> tdu -cTMu xroc '04J(dtocx:c)-ioj oj QJ CL O QJ x: 4J OJ (d-H 4-I'OX>>>Ul40t4->£0J4J^ 4-1 QJ-HOJ i*-l-r4 0>'d-'-'P^^ rrj 01 cn 0) -a Tt 4J < 4-1 cn OJ o> cn cn XI HE-HOJEcn co-P-HOij Dnw ox:)-i x3 u ri > c>4-i E-t fd 0 QJ c G x: H+JX: CX)-H1444-I0J-H 0 U4J QJ-rH0l4-»-H fd 144-1 0 E fd T3 CO "—I at 1 for woul or i in 0) (di44JOcn'H0J4J4J4J01'd'HtU 4-t fd 4-) ■HQ) ^fdCCCM 0 Id O M 3 ■rH ^4 U 0 4J OJ x: 0 C 4J c OJ l-i-H34JUUcn3 > >ix: "w cx:O'Haj(03ocHcn x>4J o-heh 3 (dcn*i-iEx:-H^-H (d >i 01 x: (d u c 4J E O 0 li'4-J'a-H4J-H OJ tT'CJ»4J Cn X4-lCx:4->(dX)-H-HO0J Qjfdfl 3idDi-HcnECC -H 0 14 c x: x: 0 H 4-) M c T) 01 QJ c 0 c QJ OJ 0 4-1 tn (d ■H 4J C 0 0) 1 0) x; E •H M cn x: cn -H cn MCQJ4-IT3 Xl-PH -HC -H 1 M 4-1 Q. 4-1 4J C fd OJ 4-) c Ul P fd fdO)>C lOJfd Ui-H 73 «*-! OJ M 14 -H 0 x: 01 4-1 C > XI P rH x: uio3 cuH 0-Htnoocotfd 3 4J(d0TJrd0>i QJ l4 H 4J 4-1 01 44 a. 3 P 4J rH 3 0 4J x:a T304H cn > o m us >x: QJ m -p -H c fd -H OJ 0 cn E 0 e fd c c rH (d CP4-1 = cnoj 4HCidWfdHWT3T3 Qjidcn T3 CL4-I rH O TJ c 0) 3 14 M OJ x: 0 QJ -p a c rH.-H ao)QJOx:a 4-i3cqjqj xix: ■f^ T-t E -H Id -H x: 4J c o x: cn 3 (d Ul P -H rH .-4 fd X T3 ■HCidM HCl afdPfflUfflCu. rn Id34J^^^n4J■Hal 0 U 4J W M c 0 4J XI O OJ rH 14 -H 3 14 (d cooj - x:(d4-icn 'Hidcn4->MM Qt3cnccr'M-H x: ■H (d c M '4-IUifd(drH4-IUi3 u (0 c Id XI C M0JOUiX:H-HC(d 4-tOCW i44-tmojai33M4J 4J 01 4-1 OJ 0)(d4JrHa'dQJo 0) rH C OJ Ul •ri ''3U44Joj4JeTJ04-JCHCoa Id CJ (d TJ 4J u (d Id 4J 0 m XI OJ H H X x: > x: 4H a Id >i OJ acnfdo a- -H -HXiouic-H (d -H c cn T3 QJ ■H OJ 3 ^ 'H 0 fd OJ 4J O M OJ X a U(CJ4hO cnr-(QJ4JO-HQJfdPOI 4J M-i c -H -H 01 x: D. 4-1 M C 0 ^ cn tn o ^ > u Ul 01 TJ 4H 3 Ul fd QJ r-i 0) I" Q. jj P rt n. r- nj 0 Q> TJ 4-t u Id 0 a 0 3 -H H u a Id OJ M ^« cn a ojowloojpuixi •EEE-'Ea~ 04J OX 4H>,OflJ-HUid-ri 3 n n in in x: >, axi c fd 3 x: • D^ E -H C 3 fd fd rH X > Ul 0 -H 4-I

i >i l4M0 M-H3C cn x: -H fd 14 fo E Id fd c W -H >, 01 Id P • rH\D -r(4-tW0 4J0JC-H ■HfOidx: -fdfd Eh ^ cn-H 4-> 4-* Ul 0 H cn 4J a (0 ■H fd XI ax: OOTJQJ = tJ-H3M 0 M QJ -H U - • (d tn Tt 3 0 •Hoooiaa4-»cx:rH3PE > x: QJ 0 -H OJ C C >i OJ -H 3 C XI C rd QJ U ■HX4JUie fttOP n iJ trt m tn i4 MfdT3otcn4-ix:E ■H cn x: a E M QJ rHCPU*>l00 04J4-> 4J-HrH0J cnarHHl4-H > E«J 0) 0) Id4-l.-t >-r4.H4J 4J H e-* 3 -H > M QJ O^H C rH C H OIHOH4H(dCn3aOI— ( OJ 04J44QJUlUl 4-ix: fa3'HUi> OJ Id rH rH 4J QJ-H>34H(a-H cn EM3E014 U X3idC>QJ4-(fao 004J •H4-IX:Xl04-' -Hcnxi C XJ T3 X ■H cn 4H QJ ooJM E o\'uirt: m3 rH(ucO'4HUirHUi T3Ui4JStn4JTJC fd u x: fo 73 4J j<: H fd • QJ U 0) 144J4J 3UO013 TJ-HCcnccucno OJ 4-t T) M OJ c O-HUWMOhUU CJCr EQJ "4hO'H0IC OJUiQrH CH OJCx^na i^(dfd-H4J x: (d 3 x: QJ QJ u 4J M OJ (d x: Q) M c Cr Ul to 0jtncTJHcr'Ui-Hx:Qj4Joo>o>*HH>,(d rH 0) Ti X3 4J Q) - -4H U H C 0 3 H X1-H-HC4-) 0JT3M3CH4JT3aC(dfd QJT3 4-1 4-t Ul 4J tn c cn (d c ox: 01QJ4HQ UrHOTJ TJcn(d--0JTJ Hcn fda-H4J3-rH0J >i Id RX:UiQJ'd>i MO) 3 0) MX14J fdo)QJTJ-H4Joxj > 0 OJ >i OJ 0 tn '4-ioioi4x:'d-H-HC 4-1 C a-H TJ ■HUl oj^x:co)Qj 0TjHaj4-J 4J4-J(04J O Q> X C Ul 4-ioox:4J4Jx:uix: c cn x: >w c 10 H E OJ vD (d 0 4-1 4J 4J C 3 U 4J Cfdcn4-) "O-HAJcn 0Cli3idQJ (d-HQJ4-i C OJ 4H 01 0 QJ QJ QJ c-Huifox:rH34J >i-H cr-HC4-*x:tn4-i(d> 0 QJ 01 QJ rH ooaJx:MW4JMc ■H X CJ l4 iH C 4-) 0 -H H ■H Ul XI D> (d cn 3 4J > CT' QJ OJ -H OJC C4J 0 otTJcnMS rHfdOJ x:oui(doi C3C "'>Qjidooiocnoj3 cnOJM CLOl4'H4-) M CT (d CJ rH 4-)CUi3UtQJ0JX: cn -H -E-HO)acc M (0 T) a-H QJ -Hoioax:ut-PT3 4J H 0 O^TJ 0 -H M H x: C1.TJ X: M l4 CnrH >, PQJTJQJiOMOX-HO 3 rH CJ. > OJ tn x: 4J fd OJ 4-14-IUl C 4J T34H4JM -H QjfittJ -HrH OXl UiXMCOJidH ^ ':;J S S ° OJ XJ M 4J cn 4H o^ x:a o)4J(OQj-HMq ■H>,>4H^£ui4JrH cn OJ30J33 4-)4-> cn fd 0 0 rH rH U|rHr-|-rt4H M OTJ crOE>fd XI a34J" cooc+3 era 3rcnox;' ■p-H0'd>iCfdUi d -30 ■Hox:4-» u ^i4C H c x; 0 ^ Q) ^ < O QJ QJ T3m4-)Ul4M-HfdQJ0J T3 0 cn c c u- rH 4J -H >, --4 C -H o^H 4JM>iidx:M rHo ac -M H x:3CM uiidx:w!jiM4J-Haj o xiatj^j M 01 0 -H fd -H u > -H ox: c ajc3 CPWUIH3 The should a ments ir and conj such. F of the d biologic substant these un of a con 0) 4J --HO H 3 (U . 0 OJ -P "0 0 4JtnfOQJ4JfdE4JC 0JC4JNCn-H •4-)4-taJ-H x: iOIUiHOUi3l4^ iM0J-Hfdfd4JEEU'd0JO3OO l4 E 0 J 'OEOMC30QJU C0)0)4H flJE4-tU0JUtUi-HQtJH4H tT'-i 01 -H t 4-J > EH4-iojx;ooofd 0 B u uirH3(dHUiooc -r^ n^ OJC 4-> 0^X1 0) (d 4->QJfdC010J 0 QJ OJ 1 •H Q) • 0 (d o >-■•-» e to OJ x: o IW > E 3 TJ X3 ia tn E fd ■-i x: o o ■H M 0) OJ tn 0 Q) •-i fd 4J 3 0 x: 0 x: Si u c ^ u 0) >, tn H u iw TJ C E 1-1 C QJ -H 4J ■P x: 0 u r-( n] (11 -H M ■ H U-i tn QJ to 4J 3 E U QJ u 3 QJ -H T3 x: -M u V4 XI Ol-H •-* j^ tn > ^ Qi in u i4 tn c 0 ^j x: >i fd c 0 U x: tn c i+H 3 IH -rH tn QJ x: -H P O 0 td 3 •-i d 0 0 > d 3 j:: c ta - 3 u JJ U QJ u x: OJ 4J c 4-t XI C U in o 01 C fd en fd ^ d c Hi fd s >1 fl r-t (A fo (d x: x: 0) T) in U T3 0 J-* 4J 01 01 QJ N 0 fd TJ ■ 4-1 tJ' td u 0 13 J^ 0) QJ OJ u ■H CLJJ U N c 1-1 ^ a fd in c ■^ e Di a-H -H 01 +J c QJ fd d - in T) x: a ■- >-i in 0 u U -H U -H s-i £ 1-1 >. (d 0) rd to -H 3 x: OJ fd 0 3 01 C tn tn 4J > d fd iH H C -H -H 4J — P O J3 u P U-l a-H Ti fl -H x: OJ 0) 4J 4-1 ■rH u C E u 0) fd fd fd 3 rH TJ 0 x: 3 TJ TJ +J -H 0) >-i 0) 4J --H o C Q) rd tn 4J tn -H C Tt O H H N x: 4-1 — 4J en > 4J 4-1 OJ 3 rH E-* 0 iH rH d D U 0 -H 0 a 3 4-) Vj (U nj to c (d OJ U 3 QJ Tl 01 y-« >i X x: H U 01 Tt -H TJ c tJ > 4J U OJ 3 3 U-i 0 -H ^ tn »4-( C ax: 01 fd QJ >i 1-1 OJ tn QJ a-H ^ fd 4-1 tn tJ' OJ O TJ fd QJ 3 01 U 3 x: 0 o Q^iH -H (U Cujj - 1-. 1-1 ja a ' fd M in c c e -H E IH -P ^ -H U --I OJ XI • C CP 3 XI a) w -H w C T] CJ -H fd in TJ (d i-t in 01 XI 3 D^-H 3 c QJ fd -rH a 3 o> 5 4J iH H E 0 -H -P 4J S-l 0 (U 4-> U d QJ x: C Tt 0 4J tn 4-) -H td y-i 0 4J +J 0 X 0 > u in fd a 1-1 QJ -H x: 4-1 iH n3 -O c 4J fd ■H x: 0 3 (d 4J C -H TJ fd ty 1-1 QJ 4J >i fd a 4J rH E 0 tn tn fd >-. QJ 3 T? fd ■H G -H -H T) flJ 4J (d QJ tn (-1 4J fd CP fd 1-1 QJ u td 01 Q) 4J fd to X * ^ -rH a 01 4-t u H >,f-i rH 4J 0 , CO to 'x: QJ tji ■H - 0 tn H 4J >, ja r-{ 3 U 0 c ^ ■H 0) u tn d o 0 a d in £ tn -H Q- tn 6 0 -Q TJ tn 4J E 'H ^ in ^4 H fd TJ fd 3 a QJ 3 a H a >^ 3 V ' U 0 fd -H QJ -H (Tj (U £ tn ■H -a jj fd -H tn fd fd a^H * 0) -1 c a u 0 ^ E rH tn 0 0 OJ QJ a u x: x: O 4-1 >i O ffl 4J TJ Tt QJ O D^ 1-1 QJ 4-t QJ aja >i x: a fd fd ^ a c 4J fd 0 fd 0 Xi N -H 4-1 4-J Q) u m c u 0) )-l (U TJ fd a 0 G. -O fd 4J 0 0 e u ^ fd QJ fd QJ TJ <-{ tn x: x: --4 * QJ ■U M to 0 4-' (X O (U -H e 4-) X QJ c Q) in ja QJ 4-1 0) tn Tt 4-t -H -rH fd en 01 3 4J 4J d 0 d - H i C --1 0 OJ ■'^ TJ OJ >i4-l CL ■-t Vi C li. 3 QJ O OJ QJ -H u tn iw M QJ Oj tJ oj a> fd QJ to C Q) 4-* fd 0 tu 4J XI y-i QJ to in -H 0 QJ QJ -rH — to ■rH U OJ > 4-t in -H -H QJ cp > a u a i4 cnxJ TJ fd -H E 0 E ■H > -H U fd 0 rH u x: U QJ en u c QJ 01 x: Qt 4J C 0) i >i4J QJ u x: a 4J to to QJ c td 01 V4 M 3 XI " 0) -H ^ 4J 4-1 ■H C 0 > ^ to 1-1 m fd -H TJ fd c 4J U 01 u X 4J 4J . 01 a 0 fd x: E a O TJ T> ■U QJ O 4J Di 0 tn 0) ■'H c fd 4-1 0 E QJ 0 >i 01 0 U H c ■H in 4-1 x: 01 ^4 3 >. d tn 0) x: c 0) dJ tn -H TJ 1-1 QJ > 3 -H ■r4 QJ ja -H -P C 4J OJ x: 4J c C -H 0 u U QJ u 0 0 fd fd 0 (U in -P M x: OJ in 0) CL-H l4 IT to 0 (n tn u tn ■H x: 3 -H 3 fd 0 -H 01 TJ Tl XI fd 4-1 rH rH E 4-) > 0 -H d 4J > cu C u fd QJ in fd r-H 0 o fd tn 0) r-i 4J > -1 a-p 4-t a 0) fd 4J f-i fd 01 p v^ x: >i^ jj 1-1 Tl Iji 0 Q) x: M C QJ ax: -H > ^ x: 4J ■H -HE 3 in 4-1 QJ C 4J a >i d XI * d cu n] 4-1 ja -H u ^*^ (d ■r^ 4J a 0 V4 X 4J y-i >, 0) 1-1 -rH U c fd ■P ^ •rH -H 1-1 X» fd rH 01 0 OJ MO TJ -H Z ^ S D 0 x: TJ tn in QJ 01 -H fd QJ -H TJ x: fd 3 -H ■H O fd 3 01 -H AJ ■rH d d u 01 ■H - 01 0 S 0 TJ t-i c Q) to 4J x: 4J 0 4-10 4-' s: x: 3 ■H 4-J > 1-1 QJ U TJ fd > 4-J TJ o a jj QJ >, 4J QJ (d T) (d 4-t fd 4J fd = -P C X3 3 QJ OJ T3 4-1 x: 0 4J tP 0 -H E 4-1 x: 0 U d 3 u S-l (D tn 0 QJ 3 iH .-( QJ i^-t TJ -< > 0 ■H cr o to tl tn 3 iH in o O d 3 TJ ft Q) — T! (1) x: c — T) E 0 CPiH OJ 3 O — T3 H x: 0 QJ — - 4-1 0 QJ l-i c -— .H OJ -H fd tn 0 QJ 3 3 OJ 3 0 QJ 2 > ^ 0) t) -U E rd CM 3 0 4J 4J tn 3 in 0 H tn -p t^§ ■H cp >.4-) QJ ^ a c ■H tn c a e c (d e 0 TJ O 0 E 01 0 0 ■H 0 x: >iJ3 ■H M 0 C 0 c c u ■H x: 1-1 -H 1-1 r-i V 4-) tn fd •-< aja 0 H -H 0 = -H 4J 0 4-1 a -l-l in tji 01 -H x: i3Qjx: 0 4Jx:cr'!-i u d a >! 0 d 0 ilMTJUd'+-t-HUHTJ4J ••-< 4JdOJ 3 0 in E in O -H ■rH ^.HOOtn 3 OJcn 4J H> a H -rH OJ o TJ 0) x: -■H 0 CPQj3X:--4 t001r-)4HTJinO -rHlOXld tn x: X 0 rH x: 4J 4J d a)x:o3iotoov400dO-Hin3fdtoo 4J tn 4-1 0) QJ 3 4J iH d x: MHJx: tncuaod foxroinu ■hu d 3 0 0 0) u tO>P3>-H E-rfin 4J^HQJ--H01rH Q) u en >i u x: »4H d 4-1 0) QjiM oua'O 0)4-1 cQx:**-iujao) E in d rH -rH tn 0 d 4-' x:o- tnEdQjQJ-Hdo 4-'>4Hd'dx: 0) H ■rH r-l 4-1 0) 4-1 to --rHfSoXlXaUOJHJ O-H04J4J Cptj s-i fd u uj V en u d toTtxo 0)E • TJa>tn fd QJ U SH M P C QJ ■H >iOo d Qjo)T)4-'aa>incnQj s-iojtn c 01 TJ ■H fd w fd fd x: QJ' ja-Hooo) inrHoincjQjH-ix^QJOi qj fd x: -rH yn 4J Q a p 4-1 x: in^Dinx:in 3d i-(MW4-ix3>4h4J E 4-1 in 'H d 4-1 Tiin faH3QJ0 o)tr>3Q dOfd d u «: QJ 01 tn UH QjaiiHO) ox:x:Ttx:fdto o>iO E r-l d 0 0) x: 1-1 0) O TJ IhUOM !HHinrH4J fdQJ4-tfdU>i^H fd ■rH u a QJ H fd e d OJlO 'OJ 3 QJOIX: E 4-tX o tn x: -H d fd >-Hx:oj>iE vocpx:£4Jot o-ho •-{ QJ 4-1 4J 4-1 4J otj+jx;4j3 •0J3d4-i ,-t a-) x: >-* u 4-1 O 0 O n. fd 0 cr u 34J-HdTJU ■H ddXi-H4-i-Ha U d d 4-t tp d x: 4J d C Qjo ^ QjdTtd>iO-r4fd xiafd 0) 0 0 4-) fd M -H fdx^toai-rHtoH-iQJc-rHXl-rH i-< "dfOfd U 4-1 iH QJ ■H 4-1 QJH 4JX10JUDifdUH 4JdQJTJ0lH adOrH4J4Jtn>, tn tn U OfdfdTJOJi-i QjTJfdO^QJ a^H 4-> 0) yn XI P d 0) fd -H OJ < 4JrH4J-H4-tO)TJTJQJ>^HQJX3infd = . 3fc.>0>rH J-iJ-iCHM -rH ox: 1-1 tn 'H fd QJ -H E > 0 d tn a > u 4-t M xifdOQJOiHdd 'diojin a^w >itj 4-) 4-1 o tn d 0) 4J fd fd Qj4JQj'Hus-iaooQj>in3 ^ in tn 0 4-1 o a 3 x: x: x:MH4Jua u3>oduoj4Jojoj(u fd fd a 0 u tn xj 3 HJOfdiHfd QJ ■HeJotnxiOPx;d QJ rH OJ d 0 iM fddwxiaj4J4-i U'H d 4J-rH d P QJ 0) 01 E x: J-i «4-i QJ 3M x:fdfdOJ TJrH x: ^ 0 QJ XI QJ x: tn 0) 4J o-Px:>i iy4-)4-ix:dXid rHiouTi N x: ^ J-, 4-1 0 4-1 ^ -P Qj4JiHinQo 3>-i oinoj H^HdU H fd 4-1 fd x: in O U din M4-I c^ ajoa33 x:td^H 1-1 QJ 0 UH -P >i >H OJ 0 tjifd-rtoj o'*H4-i4J3o CJ3 x: 0) 3 d x: 0 in a ■rHOJdOJ x:tj o-h -rH >io ^a u tn 0 4J in wJ-i-HrHi-i4-irHx^ fdETi>-ifddin>ifd UH TJ fd -P -H in tn u OJ (1) tnfd4-iuo 34Jin QJoioiEQJdiHiH 3 0) -H x: d 3 QJ tn u 3 -r^ *w4-iO3ETt4-)0)in CPOfdlJi-Q in x: 3 0 o UtOEQJ 030MlHtnM QJt-iHTJO = in -P TJ 'H to 0) TJ x: toQj-Hi-i - (no)-H>icnin>oj-PdOJ ^ U •-{ Ui m ---i c 0) rH 4-1 ■H>rHO>ii04J 4-ix:infd4J.rH>(d3CP fd O 0 3 0) -H TJ -H u 3 -H TJ-H Eddfdtn 4J0 ■H4-idr-)0 tn 4H 0 -H > u d 0 3 tn 4-1" fdox:idd uQji4fd03X3>' tn ■r^ s u 0 TJ fd 0) x; QJ 0)fdQJOE-H4-iQj-H ih OJ Tl tn 0) i-i d E en tn in ■H ^d>4-l 4-1 l-f 4-lEVH OJOJfOT) 4-1 a a fd u d x> 4-)lH-H QJOtnrdTJ U O0JlH>l-i 3 QJ QJ . to o 01 W 0 •H 0J4-ItJShQ)0J QJ •■HCP-4-tX: ■rH4-i XI tn x: QJ >th > M -rH .-1 dHJfdQjfdtnu -.tn>i4-tdinrH4JQj4-i-H O u d x:h-i 4J d U 4-J -H MrHC-rl lH>ltnXlU-rHQJfd o^hxi d TJ (T» fd -H 4J ■H U 0 Q fd -Q fd!HViQJlH34J3QJlH >i-H OJ U d iH -rH ^ 0 fd 4J p u c fd QJ-HMQJO-HOlHfdrHUtnOO-Hfd 3 0) P 4-1 E d fd QJ a o-PTJ QJ-P fn f-t tn 0)4-1 ao)-H d4Hy-i TJ 0 en a p 0 0) 4-1 0) ^ Uh fd 3rH ox^fd QJ -H-rHx: d aax:-H doi d dx:fdaQJiHTidx: ■P 0 u 4-t!d £4J^ i-iX3TJ4-t< fdtn4-ttn oiTJfd fd tn a fd u M-i -rH < 4-1 0) in d 0) QJ o x: > -p ■H 4-1 4-1 tn o cd fd 4J C OJ P rH QJ OJ iH 4-t ■P XI rH fd fd hh o "p O QJ d p d o a o -P ■H fd QJ in p xa tn 0) 3 TJ rH U -rH rH tn tn OJ ■H d 3 TJ O O >i QJ fd x: QJ E -P TJ 3 4-> p ux: QJ C -P • -H -H d rH 0) O P O > -rH fO 4-1 -H 4-1 0) 4J U TJ Q) d QJ Tl P rH d d QJ td 0 fd 4J p -H arH QJ 4-t X fd TJ d 01 0) QJ rH Uh E OJ fd XI d TJ 01 O 01 < TJ -H in rH 4J O 3 -P a 0 TJ O x: TJ p tn fd a QJ in > QJ -H > 4J P fd fd c x: u 0) ^ t-\ td fd 4J d tn o xa ■H 3 4J in TJ >. tJiH-l cn fd 3 x: E "p I 01 o I O 4-1 QJ I 4-1 to 4J xa I >, u I E to fd 4-1 I 0) o a in 4-1 u e 3 I in 0) -P E I >. I to QJ 0) fd ; o x: x: 4J o 4J P fd n QJ TJ : TJ tn i 0) d in — t x: fd QJ f^ 4J tn in tn TJ Tt d fd d d o fd fd -H TJ 4-1 d - >- fd Id 01 in rH E Q) 3 4-1 -H -p a u 4-) U O ■H 0) aTJ to a QJ -H tn ip p X 0 an-) TJ QJ 0) >i4J 4J CnrH Id fd TJ ja ■H OJ Id D^ tJ rH -H d 0 3 rH -H tn o QJ 4-* to d P in fd ^ 0) o > ' — 4-) P tn fN fd QJ 0) x: -H •- 4J U 4J fd rH 0) d 3 fd a 01 cr-H in 4J OJ 4J X Tt d 4-t 01 fd fd 01 d +J tp 0) -H tn P E -Q fd 0 in 3 4-1 tn ■H tn

4-) Tt tn L in U tn tn Q 1 QJ 0) I tn tn QJ ■■ tn 3 x: I fd fd 4-) I tj o 4J 4J tn I -p d - p o '■ QJ QJ -P TJ P 4J ' P O fd o y-i P ' 0) 01 ■ d xa TJ ■H -P ■ ij> tn TJ d d 0) -P O N ■.- _ >< tn rH QJ U (d > -H d P tn fd fd fd x:XJ Tt d rH OJ - fd fd tn ■H QJ TJ 4-1 -d ■ QJ d -p p fd 0) 4J c j:: to 0) ' 4-1 xa > fd 3 -H cn to CJ TJ -px: (0 3 fd x: 4J O u 4J fd d o 3 -H td tn (DO e tji+j • 13 -H IJ> O C C W D tn C U -H ro --H U > X M fl m a> 0) u x: « c -P C W ro O CU U ^ CTiT) --H rtj O^ 0) U -H m (0 (Q CJ O 1-1 3 E -P -1 cn OJ (D ' c 13 x: I o c -u I U H I JJ ^-( I -o c o I ^ o at x: u -P ffl o X -n 0) ■P E nl > x: ens ra -H -H ■H H > O ^-1 ■•-' tft-PC-POJCCtO nj O (TJ w O QJ -P Mi-iU!::c-H>c tT" O W C 0) [i, TJ 0) -rH U I ) E 0) ro ( o x: -p ; u -p en 4J > -H TJ U 0) ^ c y-i j: VI i-i p (D ja 0 0 (0 c c c -P CP c o w -n4 (0 -H OJ U -P w c to C OJ QJ M ■H -H C D 4J U' O W C OJ U fO Q) U Q) > 'O e C C 0) o o tn c u -H o 0 ■p a-H OJ (0 O -P (fi > u n -P W OJ OJ c x: vi >i 0 -p c • ^ u o m o u c TJ OJ -p o 0) > QJ -H 4-) -H W X -P fO +J OJ -P C -H U -H 0) 3 Q) -p x: > Di^w 1-) -P C OJ U-J (d -H O a OJ 3 o o C O M -• O -H -H QJ ■H a^ > > ja -P OJ OJ XJ (d -P ^ Vj (0 c U ni O Q) JZ Q) U O ■■-{ 'o -p e o -H 4J -H O 4-1 1+4 fO woo -H +J C V4 Tl 4-» -H O 3 W Q) C O u tn i ui I n] c -P E O U : -H (0 I CO -P CL ; nj nj E ■P XJ o X CT- (1) in TJ 0 c -P -P O nj -H -H j:: O 3 to X O E 0) m to ■r4 -H 01 Q c j:: to -p ■ '0 3^ O « fT3 ja c -I o -P x: o c 0 0) 3 C 4-* Ul -H O i4 C C o o tu m •H X ji; C 4J 4J fl O TJ 4-> I O (U glp U) >! 3 O O 0 w u o ja --H I 0) o C W -H O >i to 4-1 o V4 a) ra 0 > W' OJ 4J en 01 1 XI (0 4-1 c X) I 3 XI U -H -4 r •H c 0) > tn U nJ a-H c ^ C E (0 rH O I ■H C U M -H '4-t 0 0 O 0) 4J U^ O X) +J 17^ 3 XJ tT" C -H QJ C E -H U T3 -H O (0 C C XJ XI to -H fO -H 0) QJ a > QJ 0 o X 0 1^ o to QJ M W-l i4 rH a an] at OJ ja ^*4 4J V4 XJ O Q) 0 -H O ■H 3 to >n a c o I p e o X 4J E u o n3 m C 4J XJ CMC -'4 O n3 ■p a o a e QJ 3 QJ a to 4-> tn to C 1-t >i ■H QJ to X o !:n O X P c ■ ^ ■r4 CP QJ n3 -. to .-H U 4-> TJ -H m c .-4 ja u 4-1 o 3 (0 O -H 'H O ^1 *H o to ji: -H nH to tn to Ta a 3 QJ C X O c 13 fd 0) tn O -H ■H Q) Q) U XJ to x: CT* QJ to 4J C X OJ 3 fd 4J X O y-i X H H tn o o QJ ■H tn Q) J XI QJ XJ c O QJ -H X: 4J QJ T) u Eh (d > QJ V4 tj VJ CT 3 QJ 0) > to a O to 3 QJ 1^ QJ ja a o 4J O U XJ o a a 3 E O -H XJ -H QJ ■H OJ QJ 'H ja cnx: o (d rd 4J QJ V4 cp a -4 C to to 0) nj a QJ -. X c COO) w O 3 C j:: ■-« E O OJ fd e 4-1 4-> C O 0 tn -H u O -H 4-) fO 4-> O to H 4J fd O > J-1 -rH (d QJ < 0 C -XJ ■P >iU tP (d CP 0) 4-1 ra •-* Ui X C 'H 0 to u ■H 1-1 U n3 QJ QJ 4J 1-C Q) aE X > e to nj 0 OHO H QJ O -H S-i 0 1-1 4J QJ > > au-. (d nJ j: l4 QJ XJ a QJ C 4-> ra XI trj fd 4-1 to +J -H 4-1 rH 'H XI >i 4-1 -H rH 4J O ■H 1^ 4h to 4J ■ 4J to 3 4h -H , ■H to to QJ 4J '■ Ti O c tn - ■ XJ U Q) 0 QJ XI X) en tn c 4-1 4-t -H 4J .-H QJ C -H O -H 4J I C 3 4-t -H QJ e (d '1 -HOfdlnXCOCEC- ox-HQj+j-Hoo^4; atoiH4J x:'HUQji a c X 4-1 to 4-> 1 4JtnOQJ4-t tOtJC: to M V-l -H X 3 -H M fduato34JU4H I •-i Q a- -H to -H QJ . fd OJ QJ 3 H 4-1 X (d QJ E U T3 C -P I X QJ -H C QJ : t04JXicr>fd '(d-HUHi < 0) XJ QJ 0 0- 3 QJ O J O X U I 3 4-) td I >i QJ J XJ XJ : o c 0 IH 4-' 0 4h c 0 to -r4 c 4-» 0 0 ■H < 4J rd 0 fd ■H M .H OJ a -u E QJ M fc X XJ 4J OJ ■H to 3 0 Q to 0 4J iH c cu 0) E QJ QJ X QJ 4-> In D- 4-1 < 0 rH Q td ■H c x: 0 to ■H c: 4-) 0 OJ ■H c 4-> tH fO QJ .H 4-1 01 c a: M X 4-1 4-' W to 10 0 QJ XJ U 13 o T3 4J x: XJ OJ iH o o QJ I 4J -- > >i 3 ja ' QJ ja " XJ +J j:: QJ OJ O jJ Cn-H (d iH X -r4 (d 3 p- at U 4J -4 C O 4J 4J 3 to Q) Id o -- .x: to U -H 10 ■H XJ < (d . 1 XI ' o U tJi QJ Sh -rH o c > a-p en E C to V4 • ■H (d QJ - u a --H 10 [ • 3 QJ 01 C : cr > XI -' ) QJ IH C J 1-1 Q) X) ( to 4-J -H C to O 0) 0) ■H > X 4-» IH 4J fd nJ -H X 3 H to O >. C 3 to 4J C <-< --4 to H XJ O QJ-HQJ4-10 C3X3 C > fd U to (d QJ a QJ 3 O X ■ o a-p a4H - 0 O QJ fd 0) I a X3 e i4 J to 3 C O OJ ' 3 to -H U Id OJ --H QJ U 14 O rH X OJ X3 P 4J fd XJ (d H ■ o a 3 C to O (d (d 4J o XI c OJ — I rH * * QJ JJ . 3 rH QJ -. to O > I QJ M 1-1 I iH 4J 3 ^ a 3 ■ QJ O O X arH O O 4J OJ u ■rt H 3 u u Hu QJ 01 QjlrH '- aax (d o m to 4-t|o a c XI o (d c a o X C 14 O D> 3 X3 -H O O (d X ia -iJ X 4-> xa 4-1 QJ ih H -H a to OJ 1 > Id 0 1 OJ QJ -H Id ^ 0 to o to QJ 3 4-13 3 X QJ to l4 a 4-t C C E 4J X 4J rH rH >i ■H QJ >i >i 01 0 3 cr td i 14 0 4-) QJ XJ (d QJ 01 XI fd fd M 4J 4H j: XJ c c c 4J CO Id X E rH QJX td QJ OJ 4-t l-l X a M Oi fd fd -rl 4J rH fd Id 0 Id c QJ U4-) rH4J3x:xo QJ x: CO 4.) U 3 XJ +J 3 rl N • ox >i0-H cnoj c 4J 4-1 X X W QJ X -H QJ CO X CO 14 0 QJ QJ 4-t H t^ u 4-t XI 3 c cu M in 3 4J -H 4J 4h HH X to • OJ U 3 QJ 3 > XJ a c M tn -rH c XJ 4-1 ^ 0) rl O ■rl 4-' fd XI iH 3t-.x: ■rH 3 0 Id U o Ccn4J^>QjOiO C QJ to tn X3 Xi 3 4-1 QJ XJ U5 4-1 C 4J rH UiQ XJ ■rl C U 0 -H X X -H fd iH QJ QJ td QJ C >. 14 -rH td 14 fd O (d O 4H )H 3 ■rl QJ 5 ,-H 4J 4J 4J a-H X ■!-> >i W U OJ XI 0 4H .H 0 l4 -H c c 0 O QJ rH c ^ ■r-.4-t (d C M 14 CO tn fd c -H ■H »JH to -rl • 4h 4J l4 -H X 0 0 fd xa u 4-1 4H c 0 QJ -P -H OJ E O > u to -H fd to 4J Id QJ 14 14 4J c ■H 4J 3 01 -H td 0 rH 4-1 C c ^ ■H XJ w OJ tn 4-1 > QJ C -H 4J ■P O QJ ■H tn 14 CO £ 4-1 X 14 to C 3 ■rl E to 4-' (d CO -H fd Id U OJ O H rH to X C -rl QJ 4J O 4J 4-t QJ CO -H 0 E Id fd td OJ >i to to C -O rH 0 fd ^ +J -rt -0 > XJ i-i U X -Q ^^ fd QJXU E (d 0 4J 4J»J 01 ■H O H M C 0) j:: rH (d 01 fd 14 arH Id 0 0 c E XI XJ en >t-H jn 4J fd X3 01 £ 14 3 X a XI c XI 3 U 3 0 XJ E 4-J C X3 ■M 3 XI QJ X OJ Ti X QJ 0 a C 0 C E 0 3 QJ 4J l4 0 5 4-> fd fd CO QJ 4J •-* (d 4-J to X c li X < C 0 I "ii fd a X3 QJ C (d rH 4H CO QJ 0 tn 3 •H to to fd fd OJ -H 0 C X 4-J O X: XI QJ rH -r( Id X -u X QJ 0 «H 3 01 X E CPXJ 0 XT' QJ ■P c X QJ axJ 4-t C XJ -rl 4J 0 CPX 0 U c en 01 4-1 C rH QJ C c tn E 0 U XJ 3 C fd -H 3 XI >i 0> 3 Cn to C XJ 0 C 4J -H QJ -H 3 X ■H QJ 1 l4 -rt 3 fd XI > QJ l4 QJ C to 3 >i 0 XJ ■rl 3 QJ E 4-1 0 4J 0 rH JJ C 01 4J to to '- QJ fd 0 --- ■!-> fd U -H 10 to 4J to rd to XJ 4J 0 -H X 4-1 Id Id 0 C fd jj m •> j-( 4-1 l4 rn 4J to fd Ji 4J M ■H CO 3 -rl c tn 0 tn c X 4-1 C c > l4 -H — ' >i ■H fd a—' -H CO rH QJ Id to u Id 4-t 3 ^ -P ■rl •-\ -H >,3 cn Id i4 0 XI 3-0 E OJ a QJ X ■H Q •rH -H 0 u u tn — iH a c ^ O £ OJ QJ C 0 cr 0 c 14 O to 4-> X X3 ^ c to c c 0 X 0 XJ arH XI p to C CO -H XI QJ 4J -P -H OJ QJ >i JJ 0) Id ■H ■rt 0 QJ QJ -ri at a fd QJ "^ 5 J 0 fd iH c a c rH QJ cy x; u - 01 XI u XJ - to 4-t fd C 4-) >i 0 c ■H XI X U QJ c x: 0) tn > tP 4-1 ■rl C 14 4H (d to 01 to QJ O -H rH coo CO 4J -H OJ -H O 4-1 QJ QJ 4-» O O 0 (0 o Id QJ 0 XI M 4J iH 3 tl O ■rt c a -rf -H CO 0 l4 4-1 X3U-HMXlXJi 3 3 to M fd Id CO X C E 4h 4-) XJ 4H fd 4H 4J > 14 0 Id o -P 0 C XJ 0 ■rH 01 C == 2 ^ OJ 4-t 01 g XI C ,H g-i 4J 0 0 a c 0 4J ■H XI OJ N --4 H 3 Id o X -a -p 0 14 X 0 u > 0 QJ 4-1 QJ 3 3 -H 4J C 14 O 0 4J U to QJ X ■H X at rH XI 14 3 c-^ 01 4J to l4 4h C U M QJ -H 0 tn 0 01 a i4 a QJ tn -rl 4J C 3 rH ■H 4J a Q> -rH 10 4J <-* 3 0 0 ■H CO U 3 rH to X) a 3 c (44 3 UH Id axi fd 0 to x: to c 4J fd 0 0 QJ 3 to X a rH 0 a 0 U H 4h en CO E Id X 4J X C H C -H M Id tn CO X cr fd 4J 4J X n3 3 4-> fd c M QJ (0 4J 0 u c Id 14 4J to 0 01 e X O QJ CO XJ 4J QJ -H o ■rl -H QJ -H 0 0 to ■H C 0 0 X ■rl to ri -H QJ E QJ 4-1 4J l4 QJ QJ 1^ 1-4 4J 3 14 3 0) XJ -H tn QJ XJ 0) c ■rH 01 in c 01 4H 4J • XI 0 X C CO QJ fd en QJ 4J rH 4-1 H to XJ 0 O P QJ 0 -H fd IQ U -P >i ■'-' O 01 0 01 4J -H TD C -P w Id 3 td E U QJ ■ri ■rt 4J 0 H XI -H l4 XJ C QJ to ^ c ■rH > a '-^ Xa -rl iH — QJ H -H 4J 3 0 4J £ QJ X 4-t c 0 4J oc UfdcnaiaiQJ 4-t -rl 0 "T p a ir> XJ to i4 to cr 3 'H 0 > E-. to fd tp Id u. QJ C 4Jfdl4l4XCXJJiE O 4J k XJ O 0 ■rH 0 O QJ 01 O 0 -d 14 3 c to X at fd OJ 3 -rl ■rH -H XJ 14 0 a rH U > a > XJ 4J rH fd QJ rH ■rH 4J > C X ^ 0 -P rH CTi to E w C fd CO rH C Id QJ 3 Q) a 0 0 0) 14 fd c a QJ X Id 4J Ti c 4-1 QJ 4J 4h 0) 0 j:; a i4 1^ X fd C 01 X X c 4-t > fd c 4H O ■H ■H QJ 01 01 X 3 CO £ 0 ■H ^ C "44 X 3 -P TJ a a4J X -rl e 0) -u Id 0 OJ -H (d o u to 4H S M E 3 < QJ 4-1 ra I ■P -U 1-. ■H fd 3 SCO U Ifi ui (U at 0) +J k: rH * 3 QJ fo o fo -a -P -H 2 O 0) O t*-! ja o w O C C-l O 0 M QJ -H -H O 3 -(J -M j:: iH (Tj (Tj to (d > > 'H > Li ij "w OJ 0) TJ tn tf) W C C C nJ G ro x: 3 4-t O to M -H t-i nj O +J o a* to iM - e c — -H i^ c z (U o u 0) 4-) -H 3 x: to 3 m (u a ^ tn W ja W tJil-l V^ (d -H 0 iDt3(dUC^0fd I O Q > -H ^3 -P -H D ' M D^ x: -P Oi ,C OJ -P 0 tji O 0) H x: 3 .H . -H (d U-l ■p o o OJ (d fo XI th o Jh rH (n • *i-i td XI C -P fd 0) o (d m i^ +j IH C M TJ OJ C ^WSOCCD'Ott) ,4J4J 0\D 1-1 tU-rH ui-iC'OXi'dM-iu (u (d ^ c fd -H ■r-> Qj (d m S-i *!-( C W XJ ^ O >4-i oaj-H'HCw>p u -P (d fd --H OJ w rd CU O U -tJ C TJ -H ^ -rH C fd 'H c vj 4J w o T) x: td a -P >i-H QJ 3 01 0 -M x; p -P T) c;i-iM(i3(d --H o a ja -rH 73 > ■H a -p 0) -H u i to x: 13 c xJ 01 -H W 4-) 0) nJ Tj C XI (d o « M c O fd en w -H (d o £i C C CU 1-1 0) o 0Hl-.ird(ltfdCC'0'H ■H ja M -H -H ,4J M W 3 -^ 4-1 Q o i-H 4-1 in C CD C C OJ 4J 01 C 0) cp C 3 -H ^ H X: 73 (d CT'-H fd 4-1 Q "J e xi 0 -H • w x: c 3 13 -MOOfdOJOM 4J W -H 4J E C (d (dPQx: QJOjii'U J- 4-'ui 0 STJ-hu) c ■P>-li4>Cy-i+Jx:a)H3C'-iOQjin r--H= MJ-i Ofl-HLMW iHtji 0 x: xiafl od) Ol-iliLiU •a3X>CMx; tUrHociaii-ia) {UfdrdaoJ tno axlfduin x: (d-rHs (n4-iaj'Haj(i)(D4-» -in Qj^ C OmOHXlLi Efd 3QJ (itx:u fdWQj C4-10S-I 'LiUiD'O OJJJOl 0 c c fd w a--! w fH w > qja; iun3'w[nDa)(d>ii-i x:tn cnM o 04-ix:w0 -Hl^(lt4-l en 3 (iJ4-i-Hx:c4Jo 0 fdot wELj4Jpot/i4-ioc: Mxl+Jfd fdfd -iHoi (d jj o c a c ^ a c x: o u 4-1 = U -H fd 01 (H Id 3 ^J 4-1 t >H x: M tn fd tn rH 3 4J (u > (d fd i4 4-1 XJ « 01 01 u --( 3 E 4-t M 3 U O P fd O in w QJ 4J x: -. -H ^ C « — -. -, QJ Ti x: < tn4J4-lQt4JQJ.-H CP>i fO 'a4-iTHc:winx:fda)3'H qjc i-iinQ)Qjc:Q)4Jx:x:oQjx:fd -HQJ E0> 34-ix:>-P XICTiUIOIOShOI 4J-H 4J 4-icri-H4-i fdXitn'n'H4-i4-ic 3 fdEx: QJCfdfdOQJ QJWW4-1-H O'Ofd .H -H x: ctnLi'04-i3 4-iQjmu +J>iO OJQJ ^*(dij4-l-H r^ -ri 4-iMcuqx: c^-i c;-P4J 'C^SKS-Pmojiw HUtO=-HOOQJO ■•-i C O tu-HC u x: CM oin h 3 3 -P 4J w >i^ fd d a l-iaOfdC .H QtOE4-l oocx:o oixjw-H'Hoo uaj<:4Juxifd-H U4Juc 0) (d QJ QJ OJ c QJ 1-1 x: xa U (N 0 ^ (d 4J C T) -H ^ -H ■H 1 • QJ w E >i V4 ■H 3 ^0) C71 • tn QJ 0) td tn 0 4-1 U U fd en 3 0 x: x: QJ 3 tn fd -H a. QJ 0 4-1 +J 4-1 T3 -H 0 a u 0) fd -H 0 4-1 E 4-> c c ■H TJ 0) M 3 Li ■H (d in 0 OJ 13 0) tn 0 QJ td L4 T) 3 tn U U ^X: 4J 4-- tn QJ QJ C (d C •H tN P 0 QJ 4-1 x: (d 0 0 in QJ . ■P >i E XJ QJ -1 fd XJ X QJ ^ 3 4J tn (d ^ 0) 4-1 QJ X5 QJ W TJ fd QJ QJ r-l Lj 0 T3 4J TJ W C Ti Cr> tn QJ td tl) -O -H (C (d -H fd TJ C 4-1 QJrH 3 o a 1-1 T) -H 0 ^4 X: 3 T3 - 3 QJ QJ fd •H 0 ■P 0 0 C x: M C JJ S 4J tn d x: vi cd in QJ 0 td tn in a »P to ■H OJ QJ fd 3 3 O >i QJ 4H >1 in XI x: 0 u in in ^^ i-i rH ^ fd H o tn QJ Q) (d 3 •■ rH td p -H QJ )-l 4-1 > 4J T3 3 e QJ u c T3 ja QJ td u ■H 4-1 Q) tn • -H x: g en 0 3 4J Q) TJ QJ TJ 3 -H 01 -n CT Q) tn 4-1 x: ^ TJ x: ^ QJ 4J P C W >-( >i 0 4-) Li C 4-1 3 tn in fd 0 o tn 0 3 0 -H QJ E o 4-1 E 0 G 4-1 3 0 0 x; QJ -H 0 u 0 fd 4-1 4-1 W ^ 4-1 C 0 01 ■H x: QJ ■a u m 0 C 4J tn 4J x: tn QJ x; c 0 QJ a 0 c tn 4J -H o a td 4-1 3 ■H TJ to 3 >i c td tn QJ to QJ QJ U fd C Li 01 u p- tn - in TJ U in tn -H QJ U Di ^ Oi QJ 4J 3 C O ■H x; 01 td c x: Li u td TJ 0 -H 4J 4-1 P QJ -r-l 4J (d tn Oj c 4J td fd QJ fd tPT) a ■H X U QJ QJ P p a fd c 4J TJ QJ QJ x: 4-1 tn a fd fd c td x: 4J U 4J x: 4-1 4J x: -H 4-1 QJ >-.4J QJ 4J 3 4-1 tn c tn 4-" fd x: ^ 3 C P C XJ •H P O ^TJ H ^ 0 M p fd 2 -H rH IH T) 3 cn 0 TJ ' 4-1 ^ (d 0) T) 4-* O C i-l ■H 01 tn 1-j QJ 01 3 fd 01 i-i M fo td tn x: QJ XI 01 x: j^ u XI x: 3 4-1 XI fd in 4-1 0) tn f^ x: r- 4-1 .-H CTl 0 td p H XI C 0 p 0 4-1 OJ 4-1 m -H x: 0 td 4-1 tn T) 4-1 td ^ 0) 0 4-1 x: c td ^ c 0 U P 0 'H td < 3 01 en fd 01 QJ - 4-1 4-1 . QJ O in O t^ 4-1 4-> c P r-- fd x: C H ft a^ ^ tn 01 M O -H e p ^ -v. ^ OJ 01 fd u-l XI 01 s: E 0 4J fd p p E td 4J CP.p fO 4-1 x: tn td OJ Id Q- CT^ tn u 4J CO OJ -H OJ tn (d c -a tn 1 TJ C C 4-J p C tji 01 fd c x: c ■H 4J P { p " QJ QJ IJiTJ C td td P TJ -P fd o x: o c 3 3 0)0 0 x: c x: ■ I 4J -H 4J T3 O 4-1 nH * Eh 01 C Id C -H W TJ 0) 0 ^ -1 QJ Lo -H fd a QJ S 4-1 -. : 4-1 c QJ ■ ) 0 O 4-1 H -H fd , ^ C 4-1 -P . ^ 0 C P > -H 01 CL1 ) 4-1 > O 4 Id c p -^ ^ - ^ L-H o a o u u o td 3 U a C -H 01 -H J oi fd p en 4-1 fd : OJ 01 01 u td p ]Kx:pxipwco > 4-1 QJ (0 -H p .-H H 01 x: o 4J x: QJ Uh I X: * 4-1 to C 4-1 4-1 ^ 4-) O 0 rH < c Tl r P QJ T3 ^ o P c '^ 1 4-1 OJ fd r 0QJC4-I •■00fdC3 x:o tnxj-ppofd C 4J o ■H 4-1 4J 0 4-t 01 fd C Tl QJ OJ C W > OJ C CUO O Q^-H U fd 4J ) fd ^ td 1 C 3 4-1 1 O T) C < -H QJ < J 4J x: fd 0 4-1 1 C c/l O ■P > P C T) • C 4J ( in o in c 4-) u fd 0) QJ e x: QJ ■ 01 4J x: > td QJ P ; x: a fd I 4J w x: Id c 4-1 > . o tn o OJ p fd T) U QJ p o x: c c c tn td 4J 0 td QJ QJ C a td 4-1 ^ P O 0) OJ tn fd 01 o 4-1 u 4J 4-1 (ft fl 4-1 >i OJ TJ fd -H . . „ O-HQIP C-naOJ 3 -HO QJ ■pxi fd , QjQjaotn CQi4Jxi4-ix:-pea3tn otnx: oHfdfd '■*■ ■H 0) Cr* P QJ -H X 01 P -H 01 4-1 P 01 ■H a E x: 0 -a > 4-1 P C O P ocoj-natdpo P O en QJ QJ 0-4-1 ^ a-Hfd pua— u to 4J 3 QJ o td 01 I x: fd 4-1 > O OJ P XI 0) P 01 O T3 ' 3 0) O c x: 4J to c fd x: C 3 fd ' ' QJ J U - QJ x: c c • p m -H 4J fd 01 TJ 0 -H 4-1 -H 01 p QJ 73 P 3 dJ 01 tn td 4-1 iM 01 01 < a >i o OJ 3 -H P U en li c tn I 4J C C P P td . .. a c ■ 0) 01 o O -H 0 TJ U 4-1 ■H 01 01 fd o > ....._ >01T3>H3QJ eiT'4J_JHUJW-H 0)-Htdfd4-ix;Qjatcin ctoooj ^TJ c 4JX)E-H0)0)\ax:p QJ P003 I4J3PT1E -H PQJid-H4-i04JCUlCr C-H TJ x:x:4J cox:oQj 4-itd ojqj in4-i3td01 CT'C>CO OiTJP in ctnoj-H p-H cx:3iii 3ulCTiLi-HX:6>,(d fli-_i.i_i^ uac^ u 3-H4-I PUUUfdCeOUTlOl ouo>iOPw co-H-HCrHtn X: T3 3-HOl U 4J-H3 oioi>,OCOTlE-v QJC OP ctdxiooi o)pqjx:qj -xioj tno3 ■H0joi-HQjx:4J>tninx: M-H i:r4-ipHX3ja-*-' CO) 4J W4J0CC x:oE TJOPolo Qi:;C-H01Cn4-)pQl>,CU3CC fii jj^-r- aExitd oiQfd 4-1 T! fd -H OOIUTJE QJ0I4JO -H Ol3QJai01gC4J ^X:3lHprH0 QJ aoiotx3ac>Qj QJfdinc>OOOCTJ ■p > _ 0) > 01 01 C 'H x: c H H o > x: x: x: ex: „ ., 4-1 4J 4-* H 4J cn'O o 0 P p -H o fd CUD. a4J u e 3 O 01 -H - C 01 4J C T) 0 3 fd QJ P Id 0) -H O C X: QJ T3 01 tn p 4J x: 4-1 3 0) •H 01 4-1 fd ■-! 3 TJ 4-1 c: 3 x: u u c fd 0 4J c in QJ -H x: tn 0 -H x: 4J OJ u TJ 4-) C 0) -H td P x: QJ Tl ■P QJ -P c XI QJ fd 4-1 x: 0 rH X: 0 4-' P -H 4J -H 4-1 td 0 4J c td +J P 4-1 U 01 4-1 in c fd e 01 -H QJ C 01 QJ TJ e td g T3 01 » Ch QJ -H 0) p fd tn 0 0 CP in en M M O QJ 0 fd QJ U 0 P H TJ Q > C 0-3 QJ P C P 0) ^ 01 13 Qt O O. 01 0 x: x: -H -H c 4-1 QJ 4J 4J QJ P-H x; 3 fd x: QJ 4-1 4J 0 > 4-1 x: 0 0 en p 01 4J 01 OJ 01 -H in 01 QJ 01 CJ 4-1 QJ tn OJ x; c c 4-i OJ P 4-1 0 -H fd fd c 4-1 u en -H M Ifi C 4-1 P tn fd -H td • 0 CL OJ 0) 0 ECO ■H 3 -M QJ 44 QJ P E 0 4J ' a u p -H (ft c a CO ■H Id en p >i 0 td 0) 01 QJ C 01 -H in td a p QJ 0 -p QJ 0) X] a E o fd XI -H fO >, a 01 > P < P 0 P 4J 0) C 01 ^ C 01 0 x: 01 x: 01 td 01 c tn 4-1 in > QJ c -H 4-1 -H QJ 0 0 4J tL- 0 4-1-0 0 U -H TJ OJ 1 c •H QJ c 0 C Xl ffl 0 u c »w at Tl OJ m — 1 to ' (d -P -rH 01 -H 0 -p 'o e C U 01 -P (TJ en tn .H ^ 'C H +J a tl U 01 QJ fd -o fd -.H fd 1 0> (U C 3 (d tn rH 0 u in rd Tl at 01 -p rH Tl nj rH 0 x: M -H 0 T] ■H 0) -H 3 M OJ E 0) > cm c 3 tn 01 x: 3 ■H 4J OJ Id ^3 rH C c j:: 0 w a tn Qj 0 0) QJ Tl >i 0 0 C H 3 tji ja e Q) ^ fd c -p u j:: Qj X x: JJ £ u C rd 4-1 C 10 -H x: 0 r-i Q) >i 3 OJ 0 o M 0) -H to -H OJ 0) -P ■P -H C 0 -H o « tn -H tm-i tl rH XI tn > U U-i rd 0 ^^ 0 -p -p x: (0 4J C H U ■-H -p e 0 nJ tn fd ja OJ Cl. OI+JtJtO-HO)^>iOJ 0 u -H in ta 0) > Cr fd -H 4-1 -H -O fO J= >itn a ^1 OJ > x: ^w ■H ■p tl TI -l +J -H TJ i y-i 3 E -H td ^ >4-l cr. OJ 4J in cp o •H OJMXjetOSU-H M 0 3 e - o tl ■P OJ 4J 0 tn Id 0) 01 -H o c c nj O 0 W JJ 0 u x: U tl X .-t ij tl tw Ex: 3 «4-i QJ O >-< Q.--I ■H OJ a to 0 c T) >i<4-( O -P Q> 01-H fd -P fd 0 D> a 0 m M 3 M 0) U j:: E u rd OJ c ^ a a tn OJ tl O U 3 0 fd +J & 3 -P j: o 4J-HQJ E-PU^JTl E4-I tn E to 0 C 2 0 Q. OJ 0) 0) m • 0 avH 4J x: Q) rd OJ fd QJ ■rH 0 CJ 0 -P a in 0 w x: in u i-i (0 a Hi OJrH+J to^MHx: QJ OJ tl tl rd tl jJ -rH JJ +J 3 c (d >i u u c u ^1 fd QJ 0 U -P ^ c ^ to y-i x: 0) >i c ui >4-i M fd fd T] M -H ■P 0 • fd (d (U u "4-1 x: 0 3 u (d 4J >i -P 4J rH fd to 0 < x: u 4J M-l ^J iJ-i c «-i c r: j:: H o -P tn tn 0 0) u c C 01 n3 OJ tl 3 4-1 OJ u o ffl -rH OJ 0 -P ■P 3 cr +J in tn 0 x: ■H at rd TJ tn QJ JJ tl u tn XI o (!> O C 0) -H (d 0 'T3 U rd -P ■p cr c e -H QJ XI fd Di fd to QJ • T3 a « M 0) 0 O 4J M j:: in -H nH c fd OJ 0 0 to -H rd tl c 3 -H -H C 01 Tl E M Cn a CU C rd 01 ■p rd 0^ Id o.iDxi'Uxt'H a C rd rH U 3 ■H Tl -P 0 -P 0) M u w E rd )-i jn 0) H (d e XI -p OJ c 0 E -H Q. O U JJ tn -H -H fd u Q 0) 0 4J tJi-P tn t^ ^ - -H TJ rd ■H 0 01 E U to M 01 > JJ 4J 3 •-i ja ^ O M rH 0 a rd AJ rd -P y-i c XJ o 01 -P -H c fd 01 W x: -H C to T3 (0 0) 0 O E 3 .-H 0 c 0) x: 0 nj OJ -P -H 0 > Q 4J -P QJ 01 0 x: -a tn CL tn 0 0) rd C 3 tn D^ a CPrH Qj (d 3 x: 0) tl O > P^U -H +j M w -p e M CO u U -H , 0 c 0 0 \ 1-1 OJ -H > U ^ U 0 jc QJ e ■p 01 cr-cj fd >-t <-i -H s: -r^ T3 rH TJ (d TJ +j a xj 13 x: i-( TJ 3 0) x: O • 3 ^H •-\ ^ C O 4-1 0 III tn C rd C C cntn "C(dU3C 0 a u (0 C 0 rd IV4 Oi-H -P -H C ■H tn (0 U (d >, rd (d >. (d -H 0 fO c to -H C 0 -P W -P •H 0 E •") fd u-i rd 03 Q) 3 ■H c n-i iM c 0) x: in ^ x: ■^ ,-t fd Tl u to fd x: -H u ■H u u « tP - rd 01 0 fd fd x: 3 0) TJ y-i 3 tn x: QJ tn tl c X x: -p u th -H en C tn -H ■H tl Oi-H -H P 4-) T) U 0 -H 0) OJ u u .-H E tn Di-H in C 01 tn c 0 ■P tn >JQJ ui c U o i^ QJ ^H P l^ JZ in 0) (d 0 fN vi 'd c tn 0 O < ^ TJ cr- dJ tn 0 4-( aja tn ri] -P (d -H tl x: -H Qj 4-) OJ -H ■r4 u 01 C C 0) -H a ■H a 0 to OJ 0{n(dQ>^-p>M td tl -p QJ ^ x; a c • tn 0 3 X3 tn w Tl g •-i U Q) iw U ^ OJ 3 -rH 4J rd (d tl rd U 0 OJ U-l ■H C X: (Q M w-( u to a"a cr M 01 IfJ rH i :3 -H 3 U -H f-t 0 tn C -P T3 T3 c M-l 01 o O r^ '0) 0 * fd -HOC ja >i 3 OJ Cr -P tn ---i in 0 ■H 0 n3 at -^ 01 0 Xir-i-H (d u cpx: 0) )-l ^ C -P H U-I -P ^ CU -P ■H c -p H x: >i x: -H x: -p 3 01 H (C to QJ QJ C -P to -H U -H 3 O T3 0 T} c --( 0 in t. fd w U > 3 M H to -P ro O XI tn U D a-H 1-1 a OJ ot 3 o -p a >i -P tl tl QJ 0 c -H x: fC ■H rH +j tn x) -p OJ E OJ XJ OJ -P -H C 3 to c in w 0 (d vj fd 4-1 fd 01 3 10 01 QJ Oi u O U 0 0 0) at TJ 0 xa 1^ -p ^ tl c y-i to tn a.-p D^ > M 0 c £ 'w Q> s: rd X H 0) Id M c ■H x: tl TJ C u ■H X 0 OJ -P fd < •H --H 4J 0 "4-1 4J 3 ot -p 3 x: a-H 4J to a-H (d OJ tn QJ < 0 "d X Q> c 1-1 -H j: (d 01 3 ■P - OJ tl o ■ rd -p tn at x: I x: u 3 tn I ■P rd *J in C tn QJ : to fd c u I 01 c x: o c J *J -H CU-H QJ td QJ 4J 4J r U "-H rd C ' -H OJ Q) -H QJ - T3 tl 3 OJ c QJ -P a -H 3 fd o -P x: a in 1 tn >i-P fd : M QJ QJ -H I u x: 01 Li Q 4J 3 O 0) tl 4^ x: ■ QJ C 4-> QJ Eh x: OJ xa ■p x: 4-1 3 O 01 • c ^ ^ to rH X] 01 ^ -H to rd > fd O -H 4J 01 OJ 'H rH to tl +J "l-l 0 M O = C 4-1 u <-* O a n3 TJ • C U c 01 tn 3 H O -P QJ -P -H to TJ 01 H ■P OJ -H E M o > x: fo u OJ tl Us to to tl 0) j= o xa fd QJ m x: QJ en 0 ■P tl TJ C -P OJ 01 -H C 3 4-1 4-" TJ M to in QJ tn QJ 01 u -H > > 3 td tl tl TJ 01 fd fd 01 w x: x: M QJ fd x: o ■P -H CP -P c; J3 O 4-1 < 3 0 Tl tn 0 rH c x: 3 0) 4-1 0 x: in 3 x: 4-1 ■H 0 tn in en OJ x: c M tn to H 01 ^ ■H 3 x; fd 4h O H QJ x: = tn C tn 0 TJ P a rd C C td QJ (0 0 E tl x: ■H a 4-1 fO 0 0) U 4-1 rH OJ 4h 01 tn 0 T3 01 tl 01 c ■P 0 x: 0 u tH H QJ to 4J tl rH ■H U ro TJ 0 QJ TJ o to to OJ 4J tn to ■ x: I tn 4-j I . U 4J fO tn - o ji: -H 4-" 4-* j:: OJ >; -H . >.x: tn - Id 4-1 4h in r tn to - E in vi Tl fd in tl o tr* a Id U OJ fd x: ■a4J } Xi •-* r-i -H xa 01 in rd to tn c tl o -H OJ rorH4J04->aiaid> : 4-1 T) QJ to Id x: " •H Id c tn -H c QJ jaaiCErH+JcH I 4-1 -H -H 3 td -H tn a a >iT) j: 3 M 0 4-1 01 4-1 tn QJ cr a-H jh 3 ^ C rH 01 0 0 rH ■H QJ -H TJ tl 4J 0 3 -< XI -H CP 0 ja -H tn u x: tl ro 10 C tl 01 4J Cn4J to 0 QJ Tl -H to 0 u x: V4 3 en a - ' to 4J = 3 4J O 0 4J x: 01 C 4-1 rH Id -H 4H U 3 QJ H tl UH -U H 0) O C 4J 4-1 tP tn ■H OJ in > O Id \ c — I Id >1 01 r Tl I 01 TJ 1 C C X I to I a - 01 I X 4J ja 4J 4J u x: QJ tr- c -H C E O u in I > 0) XI l4 c c c QJ -^ O H to O '■ O 4J -H r a iH 4-1 tl 0 3 Tl 0 rd 0 in x: rH 01 0 0 Q) QJ a> ;C ■H 3 fd 3 4-1 3 01 E E t. tn x: 0 C 0 •H OJ QJ rH 3 01 c to tn tn tl ^ tl <0 QJ QJ C tH 01 -H 01 u o x: x: 4J x: 0 u x: M x: fd E 4J 4J fd 4J 0 Q 4J ^ 4J fd a >H c 01 fd ., •H 4J -H 4h 1 >ix: x: • 01 0 T3 E C at u to U 0 X 4-1 +j 0) 4J TJ 4J . IH -H ■H 0 j= at tn 4J c ^ 01 -H rH x: 0 c .4H TJ > 4J -H 4J H 3 c -H QJ f-i e-H 4J c fd -o = -H C 0 to ^ U 01 - u >iH OH c TJ fd in QJ rH C IH 10 tl en >, c rH XI u u 01 to 01 fd 4J a ■H ■H Id ■H QJ 3 c td ■-^ ■•-* 'iA M 4J Q) 4-1 Id c ^ 0 E 01 Tl x: 0 td rH Id to Id 0 Id 4J X: -rH tl Tl 4J C C ■H ro ■H tn *J tu C Tt -H c 4-t fd rH O 01 TJ TJ ■H > 4H XI 4-> 0 rH 0 'H tl rd en OJ 3 to ^ > 01 01 X3 -H c a 3 E +J aTJ tn c >i 0 M rn ■H C rH in 0 01 td tn 01 0 3 0 C QJ -H x: QJ en o ■H OJ -J= 4-1 qj to u TJ XI M 3 4J TJ 01 to UH to td to j:: 4J to x: u c aja Id OJ ii: 01 td 01 4-1 4J 4-1 ■H o ^^ tu rH >, a fd p OJ 4-) tn tl 4-1 C 01 c 3 CP 4J 3 rH en ro in 4h m 4J c td QJ T3 C E C tn T3 j:: cr ■H o QJ 4H ijq C m QJ E E to 0 H c Di tl) tl rH 0) tl fn >, 0 a QJ fM > -H <0 4J -H 3 4J 10 tl to 0 U XI S c 4-1 0 E to QJ 01 01 CP in < Q) QJ to x: QJ [il 0 - 3 E QJ QJ at x: 4-1 CP OJ T3 U 3 4H ■o x; JJ o 01 rd QJ en 0 E H fd Id in 3 0 — O ■H 0 ai> fd fd 0 = C CP ■-{ r- c QJ T3 C 4J £ in TJ TJ >. c tn 4J . tn 0 H x: 0 3 tn TJ tn Id C in rH QJ 0 01 C rH ■H 4H ti 4-1 tn -H 0 0 m 3 -H 3 x: tl 01 rH x: x: QJ QJ o ■H 4J x: C 0 trj3 XI 0 4-1 x: tl -H 4J a 01 ■H CP 4J TJ x: tn 4J 0 «^^ C -P Id 4J 3 a 01 QJ iH fd x; ^ rd C 4-1 01 -ri -H Id Id c 4J >i Id rH 4H Ji tl kH H tl aTi Id 3 3 en OJ j2 a tl QJ 4J fd x: tl XI 4H 0 en fd — QJ MH cr in -p 0 ■H E M x: 3 en fd -H tw 4h Id QJ -a to 0 = 3 « o 01 QJ 4J Id rH Tl 0 u • to c 01 01 in U C B -P x: 4-> 0) tl -H >i Id c QJ C Id 4-1 QJ £ 3 m tl m Id 4J fd • >i IH Id fd 4J in 4J a o x: 0 a Id U 4-1 rH -H OJ +J c 4J 0) Id 3 a > td 0) ■H ■H 4-1 4J X E c a -a -u tn M 4h in u to tn Id tn tl rH 4J OJ ■H Id c iH >> OJ 0 c ^ - in 0 ^ 0 QJ C 4-1 4-» 0 3 = Id tn +J in Id >i4-l rH 01 tn td x: 3 3 rH 0 OJ to tl in jq 0 rH 01 -H c o :3 s: c •-* 4-1 4H T3 ^■^ ja 01 0 >i = 4-1 Id *J 0 • 0 x: 4J in 4H TJ U c to 4J 0 CO tn C E-t TJ Ji; in C en 4J 4-1 4J 4J -H -H 4-1 0 0 E ^ Id Id >i o Id Q) C fO M OJ tn QJ > x: c ■H OJ 4-1 •-i tn tl • in 4J in 0) -P +J U H x: c c 4J H x: -H a OJ 4J x: rd ^ M c u 0 s: tn c +J +J to -H Id 4-> H Tl Id tl fO H r-t 0 u 01 ■H fd OJ o E-i XI ja >. 0 a u H IH 0 4H CJ C C M QJ 01 c QJ c 0) tl to 0 X U u 0 QJ -H c ocjx:'*Hj::4J idx; Id 4-1 e Id If) N OJ fd to 4-1 CPTJ Id Id 4J Q 4-1 0 4-1 H U 4J a 0 G £ rH c y-i E ■P ■r^ C a 0 0 xj c rtJ -P H (U 3 -'^ iOOUC+JU(U dJ d) )^ M 01 -p d) -H -H -o 1^ -i^ d) d) Ti en - CO 'U c ■*-» a Id -P x: 3^(0CP C 0 0 U E -H s <~t C Vi ifi i^O IW rd 0) 0 W U-HHJJJJ njiw 0 CO in c in (d a.j= en 3 -P CN -O -H ■r^al CL4Ji-ij3CrH dJ d) dJ > +J -H d) < 3 CP 0) -P 0 c x: c TJ ^ I iw (0 CJi tn T3 -US^OJcn-HCnJ^Qj XI J-< D 0) 3 UJ 4J fd d) c c a en (d xi—QJOiJ-i mm3 m en dJ tw C OJ C > C 3 dJ coy-iTHj:;(Tj^cjcu'0 0 c] t-i oj -H -H +j +j fO x: XJ rtJ dJ j:: "M -O W -H tP 4J c Qi x: 4J «4-i H 0) c tn 0 OJ 0 x: CPO] c X— '■H^'O en ^ ■H -P c CM -HO) aoc^ fd ■P 0 0) -P (n 0 to ro fU 4J QJ 0 ^ H J^ --^ C CU ^ CO u dj 0 tn -Pen •o-HOJI) -'a 0 c m tn mcu 4-)>>cnrH ■p m d) 3 ■H 4J (0 -rH Q) T) 3 XI in > u 'ou-ico'0 i-i ■H -H m XI Q. nJ -H 0 (d S 0) 3 x: -p -H -rH ^-O +J 0 ■-< c -P +J 1-1 T) T3x:tn 0-P>itnc c W 0 -Hinaix:(d .H-HO TJ -H y-i axi 3 CP U .H C -H ■-I Q 0 a d) OCTjscnnJO-p-p Q. 3 73 x: 0 >iw c -P c a 0 d) to C m 0 -iH -H jJ Q) E at +J o (d c> a>oo(^xa C tn a CO ' CU C TJ 0 Vi 0) -H X 0 -H "J-i C "-* dJ T-iUJ +j QJ 4J d) ■h4JO'0x:j-(W'ocx; in c x: o Wnj dJWtTJ-Pn3-H en • c dJ o to •H in e in o u H en 0 tn ■r^ jj U-i 3^-11-1013 -rtJtnxl E d) a X: -rH IW oodj oj— a-Pu 4J in ■p 3 ^1 nJ tn»4-i^c-PdJE-H3x:3 dJ to d> u ■ri c^ U 0 u -^ to O 0 Pi fl -P E H -a H 3 dj dJ dj 0 u tn C X:-H •.ptld)rHd)W w c +J x: a-H M H EHiMwoacnjJCT) d) -H tn E- x: w a ■H 01 « ^ C -H C v^ a > TJ 3 Q a u -P x: 01 d) -P "3 a to ■H H dJ 3 d) -P -P c ^ 3 -H 0 ■H T) d) j: CLox:-HiHootn 0 X 0 (d c x: en tni-i4JSo Hu-H x: « w > (d -P O c (d fd tn i^ -p (d -P -HO xt a ^ d) (d -P d) o dJ -H ^ -O fd Ti -p x: i^ -p dJ x: C >.F^ -ii 4-) 1^ -P fd x: -H a Q4 a c ■-» -p X -0 u c at c -p o dJ d) dJ c Tt in x; E dJ > -H C -P 01 xa dj o dJ d) d) -P a a o x: Id Ti Ti dJ c s -p ■-I C QI 13 H in 3 fd E CO T3 O O to C d) x: ' tn ui fd x: tn x: Q) • -P to -P x: -p in c -H (d +J o 01 •■ 0 4-1 O QJ -H in -H .H * ^^ U dJ 4-t ^ = dJ M 01 > u d) tn x: 0 n-H dj x: dJ -p u to -p in en " J w (d x: J Q x: u * -P Vi X3 c -a ■H 13 Ti Id x: 0 ■-H 13 d) 3 0 ^-^ 3 c x: t4-l -H 0 3 jj to 3 3 XI dJ (d fd +J to 0 0 in c to d) dJ d) (d 0 u > W-l a 3 dJ dJ o a 3 -P C -.H O "30 •H >, c ' to C M )-4 c dji-iocO'-iad) o <-• +j a-p -p c 1-1 XI fd a (d o o fd x: 3 c x: ■H fd -p -P tn 1-* u to E to d) -P to QJ 00 13 -P -H 3 +J > fN O -H 3 u ^ CP13 M c o o ^ -p -H 3 u 3 c dj o c oj o d) x: c fd u o E -p J.; u Q) a 3 x: o r-H ■P CL^ y-i -a >> O C -H fd 4-) ■p •• XJ (d c ■PT) o fd QJ 3 3 -P 4J u u o dj at 3 "-) -P nH ^t-l JJ >4-l QJ dt = 1-1X3 CTi OTi n ■H -H OJ > (d 3 M H (d ■p o 01 a >H X x: QJ dJ in 4J o XI U OJ -P c CO x: ^ 3 H S QJ ^ W E -H 13 Q O -r^ 3 OJ 13 -P >^ 3 C 3 -H O fd XI Tt cn c e^ fd -H o x: 4-* -H C -P fd to • o E 3 C -H 4-1 t. ^ O 4J (d o c -H tn u-i o in QJ c c u tn 3 o -H 3 cr-H QJ O 4J X x: to d) fd u 4J -H x: E 3 13 4J ij tn to o 4-t dJ tn u-i c o ^4 to c at dj -H dj -H x: ^ 4-1 M 3 14-1 c 13 QJ dj dj xi 4-1 -^ u (d fd 4J d> 3 0 c x: >-i cr c O +J >H d) ■H Tj 13 <4-l 4-1 U^ U (d -H u c fd d) 'H 3 tn * in N i-j -H 13 -4 0 c to 14 ^w 01 fd ■r4 d) 4J l4 x: +J X OJ tn H o enx: 4J (d -H 4-1 o i4 fd (d fd 14 4J CL x: 4J o E U to C H 4-1 01 OJ -■r4 in l4 • d) o c E >. a td o -H X O -H 4-) S )-t y-l 4J a o (d 4J O E C 4J tn 4-1 ^4 at HMO „ tn to fd y-< o d) fd tn a c 4-l fd QJ o p ^ s: s: cr fd 4J 4-J Q) dJ c -H o 0 13 O 3 C fd -H * QJ 4J -a cr tn 4-1 fd dl c X) O M 01 -H (d C O 4-1 fd o tn dJ tn M d) x: •H x: a > -p 4J u c M 0 fd c 0 o 4J x: -H ■H y-i 4-1 QJ 3 * fd 4J 4J O 4-1 E OJ fd -H C U tn -H rH 01 O 1-1 fd E y-* 13 Qj -H C -H O XI V4 ■H 3 M nH 01 o a 3 a 1*4 X a o X H to fd fd x: x: 3 tP fO fd xi u 4J I > 13 U dJ C QJ H o x: 0) 4J 0 c dj fd TJ x: c • fd 13 >i CU dJ ■H X 4J -H dJ to fd d) ■H QJ cr 4J xa 01 C 3 Ql 13 to to to at at . 3 c dj ■ u -H xa to ^ x: ■H u d) (d tn 4-1 13 OJ > fd 13 fd to c x: OJ x: Id o 4J x: OJ -H ■p x; "H +j to G, 4-1 rH in in -H ■H dJ 3 X rH <4H M M U in rH o J>; d) in c ■H ^ -H O M dJ - C 13 -H O fd rH 4J U fd OJ X d) J3 3 0) fd ^ x: ^ fd E-f QJ dj . d) ' M = dJ : dl ^ : ^ 4-1 fd I dJ fd X3 - u x: 4J -■. X tn ■H 3 x: O 4J dJ fd cniD cp dt ■ U 4-1 CP OJ o C O 4J 4J 13 3 QJ 0 C tn 0 fd to QJ 0 4-1 to l-l in 0 ao M (N r-i en QJ <-{ x: c ■r^ dJ H d) u X dJ M 4J 3 ■P O XJ X3 dJ o o -H O O I to r ■P 4J 4J O X ^ C 3 CLr- fd dl cr> U 4J M -H • -H 13 - s: C 3 fd -H4J0CQJC rHM- 3 to 4J -H M -H < a -H o in o o li tn o d) 4-1 - C 4-1 -H fd O U-I ■H in 4-1 u ■* o u V fd X a a dJ 4-1 0 0 xa xj u <-< QJ CU d) C 13 dJ X a > fd dJ x: U fd d) X) u d) c 4-1 fd 0 d) >, 13 IH XI to i-i to C 0.4-1 4J Ot ■H 73 3 E d) fd ■H H dJ Ti r-i ■P > u 4-1 in x: H H fd o M-H<4J>fd>lH en in . fd Q) CU-rl M c OJ -a fd -H o dJ , a c 4J . E - dj fd QJ u a OM4JEtO CQJ3 UdJC'Htnj«; oato 4-1 dJ 4J 01 u m C4-'t0inrHO QJ fd c a o fd o u fd fd M E -H M d) 4-1 Q, fd X) -H 3 C X 3 QJ to 4-1 dl d) cr c • H QJ r- -■ - -Q I to E 4J dJ c o u x: -H c d) 4-1 4J d) C 4J OJ c 3 at cr tn dl to 4-1 IW 4J I 4-) XI d) 3 E «-! to 4J fd XJOOOQiindixj 3 M > dJ IH rH X) UH 4-1 M > tP 3 .H 4-1 -H fd M O • 3Mr- Xfdfdxc 0 4J01 u-HH4Jouotntn C 4J .r4 i-l IH >1 0)ucin4-idjainin MOtOfdOE OdJ 01 tn d> E c o > X o tn o fd QJ -H dj to a t dj > iH a i4 -H (d 0) u u u fd ; M >, M 4-> -m .^ I CP fd -H fd 4-1 4J IH fd 3 4-1 X C O QJ ; M q 4J dJ 1-1 4J I fd dJ Q) E n3 rH a E X -H 4J fd , ^ o QJ a M c icxitnx:fdQi4-« n3 dJ QJ dJ rH tpx: 4J X 3 dj 4-1 in 4-1 o ^ o QJ 4J - tn tn dj to 01 y-- u u XJ M c at CP tn tn QJ fd 0 E C-H-H > OJ X) fd ■H X) XI -H a 0 c dJ Q. in fd OJ fd u fd fd M 0) X M 3 0 E H d> 0) c XI Q, 0 x: 13 QJ 3 ■H u OJ XI • --H tn 4-1 M M M 0 fd u fd at 0 fd c dJ QJ XJ 4-1 OJ -H rH tn to H i-l rH dl to rH U 0 d> to 1-1 c fd C 4-1 3 ■H 0 ■H 3 X "+-1 u u 13 in 4-* 0 ■H to dJ fd u 4H -H XI 14-t .-H QJ M c x: 0 fd x: d) 4J (d u c QJ 4J a-H Q. iH 0 OJ XI -H 3 X fd -H U d) in CP QJ 01 4-1 C 4-1 C in fd QJ fd to 0 -H >, OJ en 4J QJ fd 0 13 rH 1-1 H C M 3 4J 3 rH 4-1 dJ CP •H 3 CP to tn >i in u 4-1 c at fd 4-) H C QJ -H > 4J fd ■H in 0 c to 4J dJ X 3 CP H M (d M &«« 1 ■H x: H fd 0 QJ fi 0 14-1 4-1 U XI 0 H 0 CP OJ t4H at 4J -H fd > *.p H X 0 4J M -H M C 4J ^ fd 0 fd 4-1 0) fd c «-< u a 0 > > OJ fd 01 OJ QJ d) 4J OJ n 3 -H 0 X c x:x» 0 0) t/] 4-> < H 0 x: M e >^ (U X> OJ C 4J O 4J 4-1 cr fd fd QJ H (d tT'-H QJ E c ■H (C '-H rH -rl UH dJ 4-1 3 !m 0 4J ■POiQ'-lfd-rHfdQJ XJ tn W H -P p-t ■H CLOt (OtniHiHx:e3 Cu rH 1-1 fd XI -H c fd 3 axi > -1-* ■H fd w Id ^ 0 C M U) 4J > c exiui (D3iU4-tao CQ)-HT3X: -4Jt4 dJ cu ^M iH 1h fd 01 C Q 0 0 0 0 d) >, > 1+.I -H o x: ■ dJ nj 01 3 C 4J * dJ a QJ 4-1 4-) X: 4-1 3 QJ TJ 01 fd T3 HJ u u dj4JPV4 >iairHtnd) ja x: cr td c iH -rl x: oo^Px:oi4-ioio s ^ d) C w x; c u iti -H ni - n Q) tn 4J 4J QJ 01 0 0 IH 4J a c 4-1 fd tn c e a Qj tyi 0) j^ (0 OJ Qj rl n3erHx:TJEin>cra ^ 0 IH 73 H -rl e U 0 C UH rH -rf 0 E 0) J-( 3 4J CO x: r] ■H fd c c +J(d4-lrH4-'U x: V4 4-1 s ■-< cr 3 fd 0-HrH4-'C T3CD1 = 0) O s JJ Q) O H c Cu H (0X:'OO)hQJ3 tn4-> CT* H 0 OJ U Cm cu UH MM -H - 0 fd tt) u J5 0 >i 6 c 4-1 C 3 fltnuooo'dc EC 0 4-1 0 4J d) O >l-rH E -t-' ■• o ■I-) -H en 0 S-i ■H in (0 a tn !h u +J CXUH cu • -H x: fd • c C Tl -H (d 0 C at 4J rH 4-> o ih-c tP fd 4.) 0 QJ " dJ (d tu tn rsiG tn4J4J*w-P4-ifdO dJCTJOJCD^UMO 4-1 = 0 (J "iJ 4J rH M (iXlC4J CQJ'OXI-H f^ C Q> U 0 0 fd C -H EfdCO-HfdOOi'H fd x: 1) c QJ J-i .riOQJ-PlU'HlHEH 0 > Id XI TJ QJ +-• 4J E 3 c fd fd x: -P 4J 0 oj 0 in ja QJ Q) dJV-i^riEOJri'did tn d) l4 -H ^ E 3 O 0-0QJ4-)* 4-)fd 0) fd 0) -iH ffj x: tj ^o-Pi^cnis:'^ -H ai->H 4-1 --H d) = OJ 0 QJ UrH'4H4JM010fd> 0 j:; 4-1 T) 4-1 c (dtfiojH 4-»3d -x: (d > u (d ^ QJ (-1 1^ x: >-H fd C 0) -H 4J M iH 0 u tn 0 ■P nj -C rH 3x: 0 -rH 4J rH QJ QJ tn 0) O QJ u a- > u 3 c 3 t) T) Ch^H C iJH 4J C Q)T-idJCrfO'4H3-HOX:tfl4-'tO OJOldltn fx C M 3 0 -H 4J o dj o w 0 4-) C ^ T) c Cu tr 3 u EH C 4-1 M ■rH 5 c c fd j: 3 Cn U XI 0 3 0 -H -iH q; 01 x: (0 +jaiiHrH4-'x -Hfd 0 QJ 0 Ql -^ QJ QJ " Ot 4-1 01 QJ -HO 0 tn tn tfl x: (nua,(dcsajfd4-ix: J= Cm -P C Ti iH fd cu x:Einox:QJCPO ■H rH 0) c tn 0) 4-ix:daJuaj en (d4-) C 4-1 c tn 0 --t 4-1 jn • 01 d( fd E -P > (IJ 4-1 0) cfl 1-1 0 M 0 c c 0 4-)ajiH-HE -TJtnrH 0 0 TJ Q) >fdfd4->r-<-rl4-lrH -HWfD fd > 4J O TJ ID UH (0 T) - 4-1 U QJ in ^ ji: -H ■rH d) 4J UlC -04-ld)'H CLfd tn c 4-1 tn iH C U Ifl 0 UH 4-lQJCCUCrO ■rl r~i E_- = 3 4~< tn tn 4-1 4-> >,(ux:^'dU3'aoai tn -H fd 0 fd QJ Qj fd 4J 4-1 (UUHinx:fdOQ)C4-i UH aj:: £ 0 0) 0 -P -H -r-iH ■H E CD Vi-O in 3 > u (U tj to s^HuidJCtn o U ■H Sh 0 Q) 0 x: CU TJ 3 x: - (n ^ 4J 01 rH 01 •H C U -H -H OJ Ul fO T) 0 XI 4J ^ 3 M W 1-1 dJ iH r-( 4-1 4-1 4-1 rH fd CJJCfNOlOOlO) ft-p 0 0 tn 3 +J tj 01 IH iH Ij C (U -P •rH u w a fd fd H rH QJ 4J 0 -r^ H "^ 3 d) -rH E to * > -rH -iH <4H U ■H tT* 01 ■H >i f-i 'H ^ T) (/) T3 cn 0) tn H Q) T) E fd 4-1 • tT^-rA fd 4-1 0 1 0 mum W 3 UH H XI 0 x: HCcd P C "JH = E X: H 4-1 3 dJ -H 0) 0 Q- fd u 4J QJ QJ tn ^3 4-> 3 W IT] c QJX:C dJlH330G 0 0 E H Tl 0 Mo^p-H ui £: n -P a > 3 0 fd U ■■H JJ 0) c rH *J dj tn4-''TJ>4HX!x:uo ^ -H 4-) O rH 4J >, U (0,H014-1C-H4-) in o 0 c M e 01 XJ -C e 0 3 in to T3 0 H 4J x: C 0 o 4J c fd o 3 QJ -( ^■ri fd 0 x: 0 fd -iH cn ■H l*H u TJ XI c 4J tn 0 ■p 0 0 j:: • 0 01 a fd 4j>;tn4J 4-iC-PDi 4-1 01 Xi -M X) fO 3 01 +J fQ 1-1 U (d4-Jt(l4J4-10fdQ) 3 Q.4J IH QJ 0 01 Tl 4-idJU^fd04->d)4-l E 4-> fd Eh to 0 M o Q) ■rl QJ .- fd 0) Ex) S 0) -H d "JH N QJ c: > CUU 01 M 0} QJ H 4-t 1-1 t3 Q> +) OJ x: 14H tnccuE-H ->ixiihC X: M QJ >, E 01 d) QJ 0 cu X Cr>x: QJ 3 0) Cr> M <1) 0 rH ja Ul 4J SrH-H oai^-HOfd 4-1 O TJ fd rH -H 01 4-1 tn tn jaQjfdacntfi> fd 0) > 0 o 0) C 01 dJO'd4JdJlHQj3>4H ci. 01 H 4-> fd tp o c 0 0 M M fd en (d M c TJ 0 CT' c 0 u 0 4h x:x:cj=tnaoGOcx C d) > fd c QJ 0 QJ x: j::ooc:fdM3QJfd fd c ■H ■p (0 u 0 ^4 4-i4.;3QJtng^HU-Hd) ■rH 13 QJ TJ fd iH TJ E 4-1 jju-iH-ioCLiCntnEx; s. 3 UH 0 -H 1 « -H T) « d fd 0 U) Ul 0 ■H +J ■rl c UH ji: QJ 01 tn fd M o ■H 01 fd 0) 01 >, 3 0) I -rH 1 u 01 4J TJ Tl xa 0) 4J cu Ul d QJ QJ >iO) Ux: Ol'HXl'-H Ul 0) td rH -rl td c c^ U 13 >-{ Ul 0 rH 0) u o rH 4J 3 4J 3 C 'H rH 0 o x; <-! x: 0) td (d 3 0 ^-t -ri C XI 3 -rt 14H x: c Ul ^ 0 Tl 0 0 M -H Oh fd U ■rH fd 3 M rH rH 0 4-1 0 Ul 0 ■H U ■p 01 0 4J -rH 3 fd dJ T3 01 P M r-i M o a 0] u 01 d -H Ul 01 M 01 TJ UH a M c -H 4J 01 J.; Q) CWO Ji 0 c c ■H Ul O 4-1 01 -H . o OJ Uh QJ 0) E fd Tl 0 -H x: QJ 0 1 4J -H ■-H en 01 TJ QJ O Ul o 0 TJ M a 01 en > 0 QJ TJ 01 -H a; C 13 • 4-1 M 0 c fd C 4-1 OJ •■-i -H 3 a QJ x: c M ■rl > ■f-i 3 U - QJ en UH o -px: c -H C -rl 01 u QJ x: TJ 4-) -rA Ul 4-1 0 W 4-' 01 dJ > x: 0 01 x: QJ c • Uh 4-1 UH O • ■H Ul QJ 4J x: QJ 01 4J o UH C rH -H H Q U ^ ■H 0 d) U 4-1 QJ H Tl -H 0) o fd 0) Jfi -H •-* 01 U -rH 4J a 0 M 0 at M x: M 4-1 ot U J= M XI 3 01 +J 3 u > c x: 13 rH 4-1 QJ fd > 01 x: UH 4-1 3 td -P 0 u dJ -r-i fd u a 01 fd rH 01 - c ■H -H fd Ul O -H ^ U r-i^H -P 4-1 O c 0) tr QJ <: x: X3 a fd 4-1 tj> M • U QJ 01 0 Cn4J rH 0 M 3 0 TJ -1 3 en en at en 4J 0 E 4-1 x: UH c 01 c en QJ -H 0 td Q,^ UH Ji Ul M fd U XI C 0] 'O' d 0] Tl d at TJ ■rH 0 0 en o -H QJ C -H C -H 01 ^ c 0 -H QJ Q. fd -H Ul 0) fd ^ QJ M 3 4J -H •-{ QJ I4H OJ 4-1 c ■H TJ > 0 fd U 01 0 M 4-1 QJ X: UH iH d T) QJ 4J x: 0 fd (A 0 x: iH E C C ■H -H ox: 0) 3 U QJ >, 3 tn 0 UH M QJ 0 QJ M M 4-1 01 d fd c o 4-1 td 4-1 QJ • 0 fd 4J 4-1 4-1 U a o QJ 4J x: -H Tl 0 QJ 01 -H 0) 13 td at at r^ 0 0. c ■p --i M QJ 01 -H 4J u fd 01 01 rH a >.UH >i <-{ a QJ ■P M TJ Q. UH x: " 01 c Q.-H 0 01 0 01 C 3 fd -O 4-1 01 Ul 3 rH TJ 4-) -H o td M \ M 3 4J 0 x: 3 X» fd 0 4J TJ Q) 0 4J M QJ ■H fd dJ u fd O 01 TJ 3 0) 3 >i TJ ■p fd OJ 13 Q) 0 C M Eh >-* d QJ M -H 0 fd (^ > 4-1 E -rH C £ fd 3 0 0 aT3 ja QJ c > -H > 3 QJ a rH QJ T) -rH 01 4-1 01 CL 4-1 4-1 K ■r( U J= C UH 0 a u 4J M 0 C M cr (d 3 0 E 0 M rH QJ fd 01 u = Uh 4J d) u O UH ■H (U E 0] a a fd - a <-{ c QJ >i cr u en a4J . a 0 3 01 u x: 3 4-) < d 0 fd 01 3 M -H 4J £i ■rH -rl C 0) a 0) 3 ■ri UH rH fd U) OJ d 0 d 01 x: 0 c d p u td QJ TJ c E ■H T] u T) 0 M T) X U UH Uh ^3 0 Q. £ 1 M -rH rH QJ 'o -H 01 x: 0 01 01 x: 13 > ot 01 M 01 01 > fd 0 UH -H d) QJ C 0 fd E T) 4-1 QJ d c fd 01 Tl 4J QJ -rt 4-) Q) -rl •-{ JJ QJ ■rH fd c >. Jj O " 4J -^ 4J fd fd QJ 4-1 td C -rH OJ Ul > ■rl td +J M d 4-1 13 01 Cn4J o 4-1 £ c QJ r^ TJ QJ ■H U) Ul dJ TJ M •H C X H x: x: 0) QJ 4-1 rH c 0 en 01 01 dl 01 01 ^ TJ OJ Q) XI C 0 cu QJ Q) (d a4-i > 13 C 01 d) UH H -H 01 P TJ 4J Q^ QJ 4J a < en C 01 c QJ aTJ 4J > 3 -H TJ &I x; 0 M 0 -H E U Ul d 0 XI c £ * QJ C -H 0 0 a E (d fd M TJ XI > 3 M U 4-1 0 td QJ U d 4-) Q C a\ 0 fd c M QJ l-H 'a 4-1 XI -rH TJ D.-H >i fd M fd rH td fd 01 Ul 0 U 4-1 £ x: o 0 fd fd m 3 0 cu 13 c c >i cu 0 4J P X c x: 3 x: cj x: -H h T3 >i 4-1 dJ -rH M 4-j at x: 4-1 O TJ -rl rH C = fd dJ TJ IHH d) iH td QJ -H x: 0 Tl 0) c U) rH 0 E -P X: 4-1 ■H -rl x: 4-1 TJ m QJ 4J QJ 3 0 C fd ■H U a c UH 4-) UH ji: dJ ,Q ■rl -r^ xa 0 1 c 01 >i-P u > XI 4J d 4J iH -H x: 0 -H 0 4-> E 3 4-1 X -rl 0 M - 3 0 01 Ul UH rd . U-. TJ fd OJ •-i M c fd >i fd 01 OJ fd 13 4-1 X: 4-1 -rH c 0 0 C QJ QJ £ 0 01 0 fd M Tl 4-1 TJ 0 4J 4J XI d fd d) X: Tl rH fd en cwv 01 O 4J u x: fd x: M c UH fd M c tn QJ 0 r-t Ul OJ d Q. 01 0 ■H O 4-1 c x: CO fd E fd en 0) u Ul 3 Ul 4J QJ 0 en 0 4J M \ 3 rH dl C fd 0) -H c 0) c d UH fd en m Q- 0 d d Ul < c cr-H 4J ■H " en -H O U 13 M o -rl 0 E a 0] 01 Ul < -rl O 3 QJ u c •ri o 0 fd 4J UH dJ 01 x: c C 4J (d QJ c 0 x: M QJ -H \ 0 QJ tn 0 QJ 1 - 0 01 x: d -H > -H td rH iH M -rl TJ C 0 TJ tn cn^ri ■rH -rl CU^O fd rl 01 M x: 4-1 0) U M M a Ul XI >i C Ul M OJ 4-1 0 Q) f-i 4J td •H QJ C 0 0 ^ iJ 01 E > 4-1 d --H 1 w fd 3 rH 0 rH 0 tn XI Q) M fd C M > c c (d -H Ul x: U 0 01 01 0 ■rH UH - td tn fd QJ fd C >i Q Tl XI 01 c -rt td x: fd x; r-i OJ M -H QJ c QJ fd 4-1 u u 01 M dt 0) a 0 >ix: M £ x: 0 M d 0 -P d) +J a en C rH 4J 13 at TJ w cnx: en u -H .1 ■H x: > tn £ 4J P 0) U UH 0 3 4-1 QJ QJ fd 4J c O 0) 3 0 rH 4J c TJ W QJ 4-1 C fd UH 01 TJ 4-1 M C 0 U 4-1 -H XI o 0 \ u \^ > x: U ■H 4J 3 OJ c 0 -H rl -H D ■H l-H fc fd ■H -rl fd td CJ QJ u > Tl QJ 13 C 4-1 ^ 0 fd 0 M 13 M 0 x: 4J 3 rH Ul m OJ >i'a 0 3 QJ 13 X: X: M 0) ■H en QJ 01 c ■rl M fd QJ CJ^ a4-i C 4J d 0 +J u Ul C Q TJ 01 r-i UH QJ QJ 0 x: o O X -rH UH 4J c: x: Q) fd •-{ -i-i •-{ Xi Ul c a M tn td E d rH QJ 01 ot 4-1 0 Q) < XI -rH 01 a^ Eh Tl P Q) T) UH U rt Eh •rl M XI CL < ■H 3 QJ 4J dJ 0 < 01 4J TJ a (J QJ tn fd 4J 0 U) •-< c u - 01 C QJ 3 E 4J Ul fd \ 0 TJ Q* 01 U rH 13 01 01 H O rl QJ x: 0 3 c a fd fd d) C 1 -H TJ M ■H 4-1 d) ^ P -H cu 3 X} C 4J 0) fd > u ■p cu r-H ttJ 0 QJ H M 0 QJ 01 0 Id ^ c fd >^ 3 -H Tl 13 01 Tl M 0 u H ao d 0 fd Tl M x: 0 X -H ■H cnuH x: M 3 -rl Q) 01 QJ x: 0 x: c c Q) d -rl -H 01 x: E M 0 UH M > ■H ft4-l -P QJ TJ 4J td 0 4-1 cu W XI E (d Ul a. u Ul fO td M td X) 4J Ul ■p -H a u 0 Ck W -H 4-1 C C T) QJ -rH 4J (y 4J c Q) JJ Xi (d ■HO (1) x: 4-j T3 1 x: fd XI 4-1 0 in (0 ■H -H 0 ^-l tji D4 (d 0 x: 4-1 c QJ TJ C 4J 4-1 4-1 c ■H 1 < a> 4-1 > -H 0 D X tJi CP U QJ +J -H 0) Cq OJ O TJ -H C U (d 4-1 d o -H -H - JJ w 0 a) (d C -H xi ? E TtUCQJ^Ofd QJ T) U 0 c -^ -P E U C fl3 j:: Id x: fl 4-* *w -H T)c ■HUPrHuata C (d o • (d ■H O QJ i-l - 0 4J 0) CJ^ 4-1 0 TJ O CP l-< Qjfdi-i coa xltn P in 4J M 0 rt] JJ fO M ■rH M nH XI MO-HQJJ^rH CQJ toaotoox:x4-)4Jai TJ M 0) O 0 P tfl 1-1 W -«-) 0) U) r^ < 4-JSTlrdPC-'4|i oX4-iaj tooi-H 1-. 0 fd 4J 1-1 c tn "w 0 j:: >i c j:: WO Tl Q) -H +J 0 O 4-1 X a. Q) en tp >i 1^ 1-1 fd P tn \ U W < -U PS ^ CPTl W > C 3 -H to at 0 OJ c c w E >iXi x: 0 P in to fd D^T) d 0 1 0 U • 3 fd Qj at 0 < (0 at Tl (-l>i>d-HMOX» 4-1 to TJ cr fd -H ■H (0 0) D cn W w E 0 ax: -H u t/i tn > o QJ in CLM -H x: 4-) w i-< at TJ -H -H o 01 01 x: ■H fd oj x: uuaotMpi-i-rH tn QJ 4-)4->UOQ'4-iMai3 PL^ TJ E c u ^ W (0 TJ 4-> W CnJOJs x:WOid4-i 0 > QJ C fd C 0 3 c fd OP 0) 0) . 0} - tn 04-ia 4-lQtOJ=-H cu ■H x: fd c 0 0 enx: o >. o 0 o -H tn D en 0) x: QJ -H 0) u >, c 4J to TJ -H 4-1 o 4-1 4-ioi-ii-ix;x:csht3 ■H x: x; to 4-1 <4-i (d u +J ji: ^ > H W 0 x:fd r-icoj-a^c 1-1 fd •H O -H 4J 4-1 -H > 0 4-1 +J 4-1 0 0 o — ' M fU +J 0) ■H 4J 0 -H Qj w p 0 x: (d 01 (^ c H-IU-I+JPJ-I PtnOX3 O 0 x: fd TJ 0 ^ c s 4J at u in rd QJ P 0 4J rH -H -H M O-H^CTO 'MCM QJ 4J X» 4-1 -H W 4J CU-H IW c x: 01 w i-ijaou4-i P4JU iw QJ CfdOStoPfd >i to fd r-H > Q) W 0 W fU +J 3 en -H c -^w -H to o 4-1 C u^ u otnoiQJi^ x: ^1 4-1 fd 0 ■H 0) m (T] 4J w u u (d+J^XiOMOJC 0-HOi tn>0JEXiO tn 4-1 0 0 1-1 ^H ^ T^ M C U W 0) 'H to i^x:idO£ati-i-HTi in < ■H tn i^ T) ---I -H 1-1 m ■H 4-1 QJ Qj ■H 0 (d 0 at ^ ^ (d *J -H 3 U T3 (d tji > -H -H -H fd 01 4-J in 0 -H 4-1 Q W -U -H ^ ^ a-rH x: k-l -H U TJ -H u TJ inQjEP4Jfdtno.H> 4-1 4-1 QJ C OJ 0 ■H fO -U E (d to Xt ^^ E 4-« 3 OJ 1-1 at -H fd C Pja 0-HC4JM-HTJ tn TJ 01 [lh 4-1 -U XI — fO 3 E E c fd 0) 4-1 0 Qj (d fd 4J 0 ^ fd o OS vjuojfd C O 'H E U 1 • v^ c E OJ (d rd 4-1 x: U) +J = X OJ H J-i E tn T5 c - QJ fd > ■H en > 0 TJ -H 0] TJ U iE4-' M c ■Hr-ioci-«4Jiiotnu en o u o -H c +j J-) (T3 e to 0 u < ■HXltOC TJXI-rHC 0 T3P 00-HET3C-H C Tl P 1-1 >-i in fd +j (d 0 -H >1 T) -H 4-1 OJ 'H Tl OJ MO* n-t ■H 0 4J -H 4J (d -H 0 'A-t 0 O tn a 0 0 4J M -H (0 TJ dJ in D i*-l 0) C 0> > 4J OJ 4-1 -O U fd 4-1 0 X: to to 4-" 0 -H -H •H 4J 4-1 Q, o 0 - c c 0 ■H -H x: 0 x:a>MMMooJcn 4J O x:in(dCfdCP04J4J4-) 4-) -H "^ -H C 0 TJ a - W ^^ D H o U 4J 4J H 4-lUfdO'-lCCCTJ c < 4J ojfdEctn uc fd E ^ r-i Q) U a w 0) 0 0 )^ u C C 4J c x: CL 0 at -H c QJ in-HCLi -Hi-itofdO E -H ■H £1,4-1 p at 'H -r-i u (d ■H OJ U-l O Cfd at4-itox:cfd M -QJ XMMOIO -H J-l r-t OJ 3 TJ C tn -H -p fO TJ E OJ ■H 0 OJ ■H 4J TJ U C < 4-1 -H fd l-i>4JOfdO>UTJO 0 en C QJ QJ ■u-H E c 13 O to j-< 01 0 tpx; Ti COrHfdU CEO) 0 u 0-Hfd UTJTJl-tQJtn U-i o fd en (d x: -H M -H rH -O 0 ■H -H to W u 0) -H 4-1 x: QJ -H fd P tn C XI an 4J u 0 -H -H 0 C -U M D 01 McopQJCMtoa4J . rH C Q 01 "4-1 ^ u fd i-i s x: x: -H x: m -H l-( TJ E 4-1 -iH c u. fdi4 incpOfdoo-'-im O M 0 V4 O 44 tj" TJ y-f T) c fd {fl +j (d 4-1 to 4-1 1-1 at to 4-1 tn u OOJC U-HMM E x: -H QJ P 14 P C M-l (TJ 0 u iH > patcTi04J ooj 4J o TJ (d 4-1 CLTJ tn 4J 3 o x; tn o tn ■H C O -r^ 0 CO 4-1 c: ux:oc4-ito>i T) TlM-HC CCtT" C-H M 4-1 fd x: w en 0 tH >,-P . H 0 *--H to 4J u fd >i.-t >i Ofd4-ifdtn-HOC4J*dC 4-1 tn XI 0 3 G tn c; -H rH -H (/) at w at -H >, to rH +J >, 4J -H < 4J -H fd fd 0 4-1 fd c E at -H O -H 4J rH M 1-1 C u Q at > 0 Tt4-1 >iCP4J O fd-HM OTITIC 4Jo>x:ctr C 4J -H to U C TJ -H (0 U 0 c U -H 4J P M C -rA U -H M M C C TJ 0 fd CL-rA 4-1 0 i-< x: o C M T3 0 0 J-( QJ >-< -H fd QJ C 4-1 OtQiH-HC 04-l-r4 p fd fd 'H • x; <-* -HO o E QJ TJ 0 O O M -H OJ 5 p in -i-t J3 x; (d a-d X: PUP C XI 4-1 tn 4J c 4J in 1 -a tn 3 E x; o -rH tn CL w en u t3 fd ^4 H 4J E at Hto4-i04-io) 1-1 x: 4-) fd a 4-1 o to P C M -H 0 H in 'H fd ■rH 4-1 0 4-1 -H J-t C P E -H a > 4J 0 0^ 0 u-i en 1-1 ^ a U) a c ^ -HfdcaMXJtnu 4-> TJUfdTJ inE> TJ 0 V4 fd tn c i-( at -H (d cf) 0 ■H E 0 -H 0 U 0 a fd P QJ u >i-H OJ d^tJ u-i tn 0 -h c 1-1 QJ 4-1 -H ■H Oi 1-1 E Q--H a Q ■H U (d a= E 0 E tn TJ (d fdXJtntnfdfdOfdU-H fd a in in TJ 0. in H tn 0 in E 01 4J O C -H H -H fd -H > to U 1-1 U TJ 1 O fd QJ -H fd >i a a a p i M in E to 0 tn o 0 x: -H > en u en tn > rH c c 4-1 0 01 0 w o fd E XI E M -rH --I fd Id [d 4J QJ >, Q u iH to fd tn 1^ d E C 0 rd tn 0 0 X! 4-1 H -H E -H 4J C 4-1 O 4J < -U E 0 4J U Tl c 0 fd in fd c 01 QJ 4-1 1-^ >, M fd j^ E in 01 i/> o 4J o >i4-) 0 4-1 - CP tn C U C Tt 4-J fd 0 'H QJ -H 0 0 c U 0 fd 01 TI O TJ 4-) 3 E c x: c 10 o o fd 4-1 O 4J in x: tn TJ 4J a en a c T3 tn O -rH P H P 01 >. in x: 0 x; -h tn p in x: to M 4-1 0 fd C 4J C M ■•H u U 0 M 0 O M 0) 0) -H <: -H 3 a 4J 1-1 Tl j-i 1-1 u 0 OJ OJ 1-1 'd 0 T) u > fd fd 0 4-1 0 fd 0 -H 4J 0 in 0 4J c 1-1 4-1 >i c C < QJ fd 0 QJ TJ c 4-) QJ x: c o -H >i 0 x: 4-1 3 iH fd o a 4J a4J > TJ 0 E Vh c in c c 1-t ■H 0 0 4J ■H fd fd to u u ■H >i i-i 4-1 ■H M ■H 0 c a P c c 0 E cr 0 c -H >i 0 QJ 4-1 ■H 0 4J M o tn 1-1 P 4-1 ■H C c ■H XI fd 4J O 0 0 x: 44 4J fd > 4-1 4-1 0 c ■H > c 0 0 0 M 0 4J o C >i-H M 0 u o ■H 4J 4J Qj to tn 1-1 ■H c X c QJ o en rH OJ o 0 x: M fd O ■H > u +J r-i +J X3 c t-i aTJ fd fd 0 rS 0 0 a c H tn u ■H x: 4-) fd fd 1-J ■H y-i 4J a > TD ^ >i CP4-' 0 Tt o x: 4J c •H 1-1 fd to i-( 4-1 1-1 ) rH o a o a J fd 4J fd x: o ' 4J 1-1 r-l >, (d a o E E o o fd 3 >i o a f M XI tn J o 44 0 tn o ; 4J o 4-1 Ti 3 x: C fd -H U 4J C U ■H c c P tn -H O CP O -H O 4-1 , O -H -H 3 TJ 4-1 T3 in -ii 4J in o fd - fd fd 4-1 o >, en QJ ' 4-'rHajM4-ll-lid'H P 1-t O CP ODifd -4-'C4J'j 4J QJ • o fd o fd -■ PS 4-) in M c tn fd c QJ oi in •. c -I QJ x: o x; -H -H c ; ■H x: 4-> H 3 tn (d ^ fd 4-) 4-1 at 4-1 QJ 4-1 tn fd tn -. fd U ( X4 QJ -H rH >,£ O J t/1 O -H 3 01 1-1 4-1 i 4-) O TJ C 3 C d H C 3 0 3 fd O Li -H o in u : 4J u 4-1 d ^ d -H o fd O 01 -H > tn x: 4-1 4J d o 4-1 u > u toofd-y> -od M+JEOMoin H Wfd-HCfdrHl-llH QE4-1 x;-H(d0'0 -H in x: x: o 4H ^ O 4-1 o aTi 3 >i 3 rH XI • fd U Tl o tn QJ 4J 3 E M fd QJ o x: 4H rH 4-1 4J O XJ rH O fd QJ to iH TJ E a o 3 U XI rH o en u 4-1 C 4J d ■H O 0 "■4H x: u •-* <-{ 4J r ■ o-Hinc;in3UU in O 4-1 fd 3 fd OJ O fd T3 M d X] M fd w d U M 4J to o o a -H cuu o E a, £x o d -H fd x: o 4J x: TJ 4-1 a ^ T) O >i 3 44 (d fd o o o >i OTt4-JEHfdfdfdV4 ^ 3 3 TJ 4-1 O 10 3 C • 3 a 1 d x: o c 0 4J a = Mfdi::4-10JrH4JdrH OO-Hin rHfdOrH ■H o d -H x: 3 -H rHTIOJ CTifdlH4J4-i3 V4 d x: CP M fdid4J3V4 incM O 3 fd -H rH A-i -H . 4-1 4H x: fd 4-1 tn 3 OH d ■H Xl 4J O in 0 o 'H tn E ^ to P M to C U 4J fd to OJ £ H OJ 4-1 O in d -H 4-1 o a M 4J X O O O a a M a o 3 fd > to = o x: >i d CLrH -H Id d 4J 1-1 H -H CP fd o fd 4J rH 1-1 iH a fd o X a u o in 4-( E H T3 -H o M 4H C TJ T3 fd QJ 4J o en rH 4-1 u -H d TJ P fd o H -H O 4-1 E iH (d x: > o -H -H 4-) tn rH iH 4-1 TJ o 'H o fd to d T) 4-J in u O M o 4-1 fd o U TJ 0 d 4-1 P o 4-1 XI V4 Id fd lO fO r-t tn Tl d M 0 a 4-1 o fd W tn E u 4-1 4-1 a -H fd 0 in M tn o o g = i x: 4-1 4J M W Ul H fd to < QJ 0 ""J s z g < D Z 3 O u z u < z g < Z .-. («_ « < o - rtj CM T3 +J .— C o *- c «o LO O +J • C 1- en o t/l IT) I/) <*- O T3 a> -r- -C o **- +-» 4_> (O tn > «*- C E T3 ■"- O QJ -I- C ■»-» e en— o o 1- 4-> tU l/l (TJ fl> S_ CC •— T3 -Q ^ UJ QJ • E O 0) ■ — ' ui t/1 5) QJ r— TJ • E ■.- t/) 10 Q) m 0) (U **- .c on o i- O ''^ 4-> O 1- *" QJ ro "-^ ■*-• C7> QJ 3 QJ C ftj ■"- (LI s_ ce q; c +J -.- o g -S "" > <*- Qi B B r— QJ CO ■'- S_ +J ..- O i/l ^ CI- rtj E e: - +^ c ^ © C 1/1 o ^J C7> O O +J ->- -i-» QJ IT) -M 1- 4_> -.- -a c iC *J _i c ■<- >>(. -O O 3 U C C T) o o u c < 3 Q..^ Li- — Ol - o e *^ **- en Li_ 1— . U Q> -^ ra ' ■>- 1/1 c o a> O ■>- QJ QJ O -t-* U QJ O •r- en ■o -o >♦- c ai o QJ .— CL 5 >, C3. O U IT) C C 1/1 ■M r- lO Ul > QJ Ol Oi -I- 1/1 irt U ■!-» 01 l/> C U 1- . E « 1- n) o . — 13 "O ■'- (a E C Ol I 1- -I- 3 O I J-> i- **- •— CO. O QJ 1_ -.- O U- O J3 I o *♦- ■ Q> -O ; ^ .— -a c ' ■*-> O) QJ m > QJ ul Qi C I— Qi •— QJ r ■ QJ •<- >f- c o ^ O CJ ■»-> .— E 0> .^ QJ +J C 3 U S- •-- •^ fO -c • M- Q. in S- »*- QJ (O E O Q 3 C ■*-> C t/1 Q.C. a> c *! ■ 3 , o E *♦- "^ +-* "C ' i/i c o en i/i c -I o • — ■.-■— -tJ 03 o c +-> C U I •r- IjJ IT) >,-i- ■■- -I -t-1 > ^ ■»-> ^ (O +J 1- fO >, O I Z 1*- QJ 3 l_) 1- ' QJ i_ C -Q > >, cr^ r— CT> QJ Q- 5- C m O QJ ■<- C QJ Q. C O en wo +J ■*-> U J _ Q) -r- : Lu -o -o E +J QJ i_ >) 1 ^ « S- QJ ^ < . U O -C CL J . QJ E *J -t ■ 4-> 3 *- ■— O 1 O -tJ QJ O (O • - 4-> I- 1- QJ -*-> C o>>>fO 3x:.c i/io C ■*-• 4J 4J ■•-> LU ■■- •^ ■.- >, 3 *-» 4-1, ■ l*~ •/)■«-> U i/i -,- j3 c q> c «a: QJ ^ -.- i_ O ■'- Q) >iTJt/lfO-'-'«-E'— i.JZ) 1/1 QJ-t-'-'- QJrtJ rOOOCO+JQJl- .C S- Q. fO 1/1 i- QJ Q. QJ 3 cma r— -O -C <— -i-O"* ro U- ■^ 4-> 3 S- >,.— UfOOC QJi — fO"D j_ .,_ (J .^ -o ^ C QJ QJ T3 QJ 3 O 1J1 E,.Q QI i- Q, *-> C QI to 'I" *-' QJ fT)'T)>Ol/lC-'-'1' ^ i_ O 1/1 O C -C +Ji/lfl)QJOlJ'— •■*-* QJ (_) C C 1/1 Ol/li — -r-l*- O 'O'O C QJ TJ ■*-> O •<- QJ IJ ■•— U fl 1*- = -OS-Oi-QJ 3 E >OQI-«-*I=UC-i- Qji/lECiOi/lOCT QI E (O 1- •'--'- QJ .,- QJ ^ .— QJ ■<- C 1/>C+-»*J S-C+J O rO ■-- (TJ <*- +J O ■<- 3 1^ -C C Ol+J l/)(0-l-)aJ(0 --QJW j_ e *J in Z > QJ l/l fO 1/1 r— = ^ Q. Ol QJ QJ QJ to QJ c "c; 1- i/i 1/1 • 1/1 ij0T-3x:inOl-C l_>r— ■t->30-30 QJ -r- u U O O O l»_ ^ c -M to i- **- 1*- o to '■" Q- QJ on +j >, ^ ^ '"^ "^ I. I L.-4->i — LlJ-i— Q-E fO .C QJ O O O QJ +j CT> > QJ QJ '-J j= C -r- QJ -C CL**- t— ro 1-.— I— t/1 OfO 3 ■<- T3 Q..C QJ -O r- lO -^ O O -£= ■*-» , j=t Qi I -4~> > ; en c ■»- c QJ -tJ I ■'- E to I 3 -M -■- (J at I O O *- QJ C --- t/1 > L- C 1- C O T) O ■<- (J OJ QJ O QJ O QJ t— : QJ -a O E C V- ■<- «0 Q. en ■t-> 01 > FO QJ O C i- s- : +J ■— >> ■ c "- 1- . t/) *o < — . 1_ Q. rtJ ) 3 -.- -r- I O U ■*-> I u ■-- c QJ QJ x: I il ^ X - I— +J QJ . 0 <-* u m w 0) +J 0 x: dj in H x: 14-1 Jj 3 U M u U T) cn (n TJ ->-i -a 0 C (d H EU ji: , (d TJ 01 C W ^ a E 0 a in X > ■H ■H i*-i P X) Q> 0 a>4-J u u V4 OJ ffl 13 to (d OJ a 1^ > ■H TJ XI 1-1 W TJ C 0 c 0) (d ro 13 4J o 10 QJ -P iH -rH c ^ > U) 3 4-1 0 -H tH 0 in 03 u OJ 4J 0 s: Ul > 3 0] Tl 01 -H in 1-1 CPtJ 6 u ^ d Q u QJ «3 ■H ^ a x: QJ in QJ m H ■p s: 4J M fd t^ 0) x: 0 c u m rH 4J +J 0 a> in a H nj H x: e 3 u ■p ■P TJ n 0 0 Ul 0) u QJ E XI (0 tji e X) CJi i (0 ■H CT' td c a Q) Ol c 0 x: QJ ■H n 4-4 c +j 1-1 ■P QJ n3 0 (0 4J 4J •w C QJ CO 3 >i 0 0 > 1) in a-H ■H U Ol ra l^ C p (d CJi CQ TJ 0 ra x: 3 ji: ■H .H ■H > CO u 3 in M £ 3 QJ U in 0 ■H 01 B tJi 3 to u x: in fl 4J U C QJ ■p fU C ^ CO 0 +J c ttJ tfl ■H u CW •H E ax) = ■H 0 m QJ 4-t -O' in fd u c n 0 3 3 u U 0 0 ■H 3 4J rd ^J jj u T3 c (0 CO QJ (d 4J -P x: 4-1 x: C 4J 0 tj> QJ c ■■-t -H U l~l QJ in x: o x: ■H c Q) TJ H to E C fd c H (d 01 0 cn-P (d H QJ in CO iH (U to -p o 3 CO c c U QJ 0) 0 to •rH E ■H M QJ C X3 QJ Ui 0 j:: fd ■H in c in ■i-i fd 0) C/1 U-l e 3 M U [d fC QJ in Q x: M "HE* XI a QJ CO 0 E CO QJ rH H ■H tn 0) tn x: in > 0) H 0) 0) u C T3 M C fd QJ 0 Q> Ji -rH 3 -P -P 3 01 (U (d c c 10 U T3 0) 0 c 0) rH 0 iH 0) x: CO u u +J H C 0 QJ C M 0 UH 4J QJ rH 0 Q) (U E -rH xa XI 0) 3 rH u 0) (d •H cu +J u ■P E E to C CJI c -H 0 CL 0) fd C c td ■H 0 rH O c 0) e u QJ ■rH 0) C C D^ fd M o 0) c -P in ■ rt rH fd fd d) 4J jh -P fd x: XI fd o in u -P -p X: yn QJ -ri u Tl -P > C QJ fd ^ C M J= E 3 H-i O nJ O -H rH < j:: Q) -P ■ (d CO Jj H C 0 UJ H E 14H 3 0 u > 4-J o Q) 0 u fd 4J s: v^ QJ x: ■p QJ fd -p ap 4J c U 4J C 3 IH rH ■p x: H QJ 0 0 u +j > HH •-t QJ -H 14H 0) fd a 3 o 0 X Uh ■-{ o 4-1 0) QJ CO HJ QJ X; c iH u 0 XI HJ ■H QJ o fd > 0 1-1 a 0 c tfl 0 4J H-l E -P Q) (U c ■H > E 4J x: o C ■H 0 c a u 0) 0 CTi O QJ fd x: CO > w 4-) -p fd OJ QJ IJ> o 0) cn E U fd ■H HJ V4 c ■rH 04 iH IH fd 0 tjl fd +j ji: 0 ■rt QJ 0 a tn 4J c 4-) U 4J >iC 0) rH O x: 4-) 4J G c 4-J £ 01 u 4J *J to fd u QJ QJ in u M •*-t ■-i H 01 c a 0 I4H r -H 0) 0) 4-] 01 Q> u u 0 cpx: CO c c 4-) u 0 fd r( 3 4J 4J 0 4J 4J 0 )-( QJ fd o XJ 0 c QJ x: QJ 0. 0 E 4J -d in E iH QJ H in 4-) c >H > Ul C 0 M QJ ^ 3 0) ■iH QJ V4 td U u CO IIH in QJ 3 QJ >i 3 H IH M rH u TJ U a rH ■rl 01 c fd 4-1 fd s: 0 OJ -H IH 4J u x: 4J fO QJ 4J c a-o »4H Q) fd 3 0 x: 4J 4-> UH ^ 4-) 0 in o U to c § c -p O ^ 4J CO m 3 ■H 01 0 to -o 0) > 4-) QJ td c X) 0 IH 0) 0 £i c/) ■H X3 fd l-H QJ 4J rH [ij x: >i 0 3 x: Q 4J fd fd 0 p e s: Vh QJ X) •-i 1/1 0 s: c x: (d l*H 4-> fd u IH m QJ M 4-' in x: TJ u 0) 0 0 3 Q) Q (0 > [1- C H fd 01 tn O 4-» -H TJ x: 4J --H fd rH 01 E-4 c in c td in QJ C iH IH o E fd 01 4J a E a4J in o O X rH 3 IH u 01 fd << a o < o I- O (A Ul (0 Q m H Z 3 eg 00 a: ^ s +j 3 G O H) 0) » £ B M nj J n ■u n oj 0) o fa " d 0 4-1 y-i m u w o ■H n3 l^ • -H 4J O a > iM d U) d o d OJ B M 5 a; ^ n) c 4.1 4J •H M-l d o (d ^ 0 a> u •H -a .-1 S OO U 01 ■H M IM m i a cu ?l en a> O r-* f > ■-t « & d o; ti) 01 en 3 4J d cS ■H j= T) 1-1 ►-t 01 (J* X d 0) (T- O CTi lu i &o d g.5 (u n 4j ^,/ c o o •r- (/) ■<- o J <+- QJ 1- O -r- o -•- C +J .— .— O ■>- -M O , V, cn CU S- ■M 1- T3 -C Q. OJ QJ -C *4- (O O +j +j a> -r- l ' Q. 4-> ■• a, +J fO ■*-> C »D "O ■■- O U "O C -.-■.-£ 3 a> -t-J -M zj O t/1 T) u o 1 M ^ UJ O Z X m o a. uj Q — _l 4 O 4 ui -o ui X ** m f^ 'Z z ■N. CD Q Ui o ff Q -J I ni m »- UI 2 in o o K» £ u. (T rv O 13 09 X z > u to ■«l z o -o X t3 3 o- •« UJ z ^ UI o Ui •-• -^ K> _# •- I o — _» u. •4 0> O »- 4 »~ 3- •-• i9 OQCCCUiO OC Z z *jou.»-i»-Q4 :* »-t z a. z X u. >■ UICLO«0X4 0»- •-•>- rjz»-«*j —I 3 _i»->-»04 at>"_ia>o z_i z4»-nuicn uuoo»n(ruit-t(E^-«4 •-•OZ t— UiUitt)^ ^ LJ I— U O <_> O UJ • i_» (rfr-crztti-LB— to Qcui»-z:30 ozoo. ■4t3COulOO*-»3«-»a. 3 i~Z x« •-•o*- o Z •«>- Z ui_IU,0»~Ui<0 4ac»- CE Q:4*-«a:uitf>uj ouiuj »-«»-a.u>a:a: I- •-• :> uio z o zuiciO >uiuiaz a'~»z»-*z x»— en uiwiu-x ujx Ui CO en to o >- »-la^ZUlUJQC ZUi>->-* -40-4*-a:r)Z OOT_ii »— iu-4Q.OO»-»»-«t3 UJ«0»J>- »-OZt3 [K 4 - juiuj-iuio:*-* UI-UiO*-«*-»«-»3X*-— * 4LJt_>uixZ4CLui(n4 Q. UI >- UI — I O UJ xaz a-tt-ua o-ujuio •-•■-«4Z4a: (Dz a(_)Z}Xtajui_i z _i i-« (-^dt-jz •»-• 4wa:uj z •-»o_i •— ouix >-oooa:»-»4X z ii-t-o»-»«»-z»n UJ04 4 TO -Zi >-* *-* Itr >'Ui»_»4Z4U-»- Z4Uiai[K*-*X04_> 4 ooi ►-•— a. o> <3L tn*- "y- tjx4»-zac •-• U4a:uj»-44ui ^ (^ 1^ •-• 4 4 UI *fi zxo-»u.»--ioa:i: z UJ»- OOZ4 t3»-«o at a. 4 •-• Ui — Ui U.(n»— 40UlUi»-»4»-Q 4uia. •-*xa--ixzz^ a:ij4ta.»-»-c»>uit-'-«4 3 Q at I o 4 »/> c 3tJ •-•U.»-4UIIZ^ OO I— •-04ljll3»-0i »-«4»-»o I a:o»-» ui»-»:3t3x»-uii3mi— -»cr *.>- zujor> «ooxui-ac ai 4uj- (LM_ t-ito^uji-i-tnr (oocjuJCLO »— a:oz> UJ44X 4»JUiUJ=>30* acx«j»~acui2 ou-Olj- z »- 4 Z (E Ui o o z >- 4 or UJ (X> U- r^ O § C 4J 4J I H O I 3 x: x: ■ rjj.;3-HCifln)Cx: n] (d o ■ E J= C ■ ■H s r U 0 a c -o M w to u c m I 0) E ■ E >, 01 u) (u a 3 - w x: > 10 M ; -P U) 0) flJ ■•-' J -u E 1-1 -d w C 3 0) (X Q> 01 0 +J 4.; TJ I e x: in c u I Q) >,-rH O -P ■ Cpx: tn +J « I 03 in - (fi x: ' c ■r^ 4J . 4-11 rt rH C N 'O ■ E XI Ot H 0) 4J ( d E > cn c : ■■ 4J 0( ffl q; ■. in ct> — C > -I 0) (0 t3 fO 0) I c a> E M I 3 o ,<4-i +J 4J ecu 1 0 0) Q) QJ ^W >i I 4-j j:: > o Ti rH M in n] 0-1 -H O H x: C XI I rd a'4-i 0 < ■ E - - ■ 1^ i^ 4J k a-H 01 -i T} QJ QJ U O to x: x: a -P 0) -H -P -P M (0 M O O -P -H 1 O -P 10 3 . E -H e Q> QJ CP ; -p M ij O M T) QJ P C T) 3 Qj cu x: x: E p E-t 'H Cn 4J 0) fO . — M x: 3 Tj oj - -I M 3 x; 3 QJ m P 4-> Cn-H U-i 0) 01 '*-' ' 01 > o tn i^ 13 u oj I (T)VjUC>H(0 U 10. " u a-H 3 -P -P fd-HoiuudJoc: :i vj o x; -H OJ U 0) C P ^ o >, (/) 01 ■■ I cu x: u 01 J M -P CPXl ^ x: « c 3 in 3 ji 0 in u c OJ 0 M u 4-1 c t-i in 3 3 0 Cn >i in c 'H x: U -P O 3 OJ O tn U OJ fd tn -H ^1 0 0) OJ OJ (U M vj XI a 0 Qj E in E x: 3 4J C 0) Tl c c 0) 01 o cd M CP 0) t4 O >1 x: (0 4J ^ 3-1 Tl d) -H tn jj CT a M 3 rd rd 0) XI E ^^ ■p fd - (d in Tl 111 in 3 M MM QJ -P O 0) M -P rt] E -P rd fd x; nj H 3 -P E 3 0 QJ a 0) o -p o -p in w +J x: rrj >, td 4J M * in M ■H OJ in o OJ 3 a 0) u a E -H ot e tr 0) u 0) d -p ot 01 -p -H a jr M c in +J c fd -H -H dj x: JJ -o c u 3 c nj rd O fd OJ x: Ot x: x: j^ +j cn+J rd C 3 CO -H O >. OJ QJ M iH S -H in x: ot U .H 4J > 0) fd -H — a 3 in +J XI Ul TJ Q) U — -H --I 3 M > M-i M Ot 'H -H 4J 3 TJ e tn I ■H I fd o ' u u ■ot OJ u u Tl > rd a to QJ U ■ u o td > O X >p Ul to Ul 01 o O -H U 3 -P > M O (d C rd 3 1 C o x: o o ■•-\ (n C QJ 0) tn o c c 2 O -H . tn ot < at x: • E 3 I QJ t x: c u fd I tn x: •- : 4J T3 ( -P CJ I c tn N I at 3 'H ; E O C 0 -a x: 3 iW QJ 4-t QJ ■P T3 x: C Id nj C Ot *j td — t H C j<; 3 M x: M 3 in u 01 QJ QJ o 0 x: fO H > U .H ■H -i-\ 4J P -H C Cn x: fd in l-i 4-t O c 3 E - u) (0 td 0 iH x: u tn Ul td -P >, >, OJ Ul QJ u T) ^ a QJ td •> U OJ td 0 M 0 -H 10 U E QJ X3 3 U ■P 10 3 0 U-. Ul c 0 (d 0 o ot >. 01 C Q) c E Ul OJ x: M M 0 tji 0 QJ XI -P 0 rd -H rd QJ -H U *J U 4J Cnxi TJ ^ fd c I-* td QJ -H o ^ p OJ td C QJ x: 3 3 0 > C td C 4-1 0 c D^ C C 0 rd 4J C --( ^ E C QJ U O O 3 > l-« H U O 0 U-l .r4 fO 4J 4J 3 -H 0 in +J fd c 4-1 3 C 3 O OJ 0 arH < 4J U E U 3 U -H -H Id T) QJ XI U -H QJ M ■H QJ M Ul OJ 01 X U] -rH to 4J 0 p E a ai 0 4J 10 td y-i U H Ul a c 0 ^ td cn x: 0 «J a 3 c U-l ot CTi 4-1 M Cn-H M C ---t a QJ in 0) OJ U H 3 x: ■r4 M 4-1 QJ J^ 4-> QJ 4-) 0 j:: td Ul T) x: 4J Tl U 4J QJ -H C H 4J M .-H a ■ 0 3 X fO 3 3 C at XI 0 *P 0 TJ < C -^ 3 O H C ot rd T3 U) QJ 4J rd x: at — QJ -^ 01 OJ Id • 4-1 U -H E U-H at Ot o U -■-i u :2 in c x: QJ cn 3 a 0) 0 4J c a ot O 0 x: H M 10 M wi a T idable both w Southe ||l|o?l3?ifs'l tf H 2 tf 5 0 W o < 2 o tn J < k1 >h < o h-t t— < O h1 o o m (1, OJ Id ■H K-i 4J c Ul QJ 01 c 4J fd fd S 0 4J u m -H iM C O O .H O It ^M P ^ £ 00 cn ■r4 01 -P r^ C SUM H •H Id E x: • ii-i a o Ul M ip ot O rd s o Q p; s QJ E 4-1 4J -H Ot < U U QJ Ul Ul 4J Ul OJ 4J O U H a o <-* u QJ Id a QJ M OJ C C M C -H M QJ M O E rd QJ E x: 1 O J r-l fd Xt ■ a u 3 E -H a ' M 13 u-i QJ TJ 4-1 P QJ O 4-t fd +J rH QJ x; td .H 4J a >, •H M Ul C u td -H o ■H 3 x: P M P Ul MX) 4J rd QJ cpx: acn c cn C M-l «-) O O M T3 ■H c at > O 4J M C 4J O at o ■--< x: P M 4-1 C P 3 J 0) C M : U QJ 01 3 J E M O _ QJ p fd ) H rd OJ O a 3 QJ C • Q TJ O tn c -H M QJ Id Ul td -H : 4J > -H > "4-1 M O Id QJ (n M Ul TJ C 3 O O fd u >i 4-1 4-1 tn > >P to M w U OJ 01 TJ x: P 4J 4J IM 4J 01 rd H x: >i M M 4-1 XI T3 3 4-1 QJ Ul x: E QJ tji at -H -H 3 P tn M O rd QJ 0 x: 4J L, P 4J CO td m -. u: ot 0) Cn-P x: u •H c P M Ul QJ 3 U-l U) QJ C OJ 01 -H 01 x: 3 > x; 4J th ip QJ Ul •-( ■-( x: -H td rH 4-1 o Ul 4J a M p fd o o c x: x: M-i QJ 4J E QJ Ul E 3 3 0) 0 0 M U C TJ rd M C Q. fd at to QJ 01 s - a _ c 4J OP 4-1 M 4-1 -r4 OJ rd fn E 4J M rd X) 4J QJ 3 01 TJ 4J Ul -H ■-! C Ul QJ ■ OJ Ul c E c S -H o -P O x: u M -H 4J fd Ul .-nam O 3 OJ QJ *J "4-1 Q W >i4-l O Ul ■H 3 O 0* CQ O OJ M 4-1 -H tl O M td 0 4-1 fd 4J TJ td a c td i-f OJ in 3 QJ E Ul 4-1 x: c fd Id 4-t o XI U ME' cn C 'H f ) O E J QJ -H C J3 4J O fd O -H 3 iw rd 01 <*-! C TJ tn OJ 4J M C td OJ U -H 2 u 3 H p c at «-i i 4J w >ix: C 4J -H 4J o «-< a-H fd E 3 P* M O C T3 U Ul 0) Ul 4-1 O XI -H C x: 01 >i O +J E 4J 4J 4-1 -H 4J M .-H 3 fd fd fd 0 x: a 3 >i-p QJ cr c o fd Ul x: Id Ul u a 10 0) QJ E to S 3 O 01 fdutntd wui-HC C iH E M 4J o iJ ^0Jid3 4-iOlU-Hi 3 -H Ul x:

icn u at ^ QJ > QJ 0) 14-1 Ul O C x: 01 3 rd U4-14J rd 3H>PrH c OJ tn 4J P C QJ TJ - 'PC Ul -"-I E 'M QJ o fd M 4^ M «-< 2 0 — I OJ l-i 3 fd 01 ■■-( "W > td O 4-1 01 >4-l 01 QJ a4J • td 4J -■ QJ 4J U S t/3 < QJ 4J Ul rd X: H 4-1 M 4J --4 OJ TJ i|-i • V4 QJ O >,■'^ ^ u Id -I o aip td -p E O > 0 H o td Ul M ip j= cn a Ul u 01 a -H -H c fd 4-1 x: TJ fd 3 c J rH in 3 ( .-t Ul 0 ■■ td >, P Ul > 0 u-i Id J o x: 01 4J . 3 TJ C M OJ ^ 3 4J 01 E 0 H > C ■H O QJ -a 4-1 M 4J C C H -H H fd > au-i 4J c Ul Ul QJ 01 ai X) Q 3 3 01 to JZ 4J P rd TJ 4J Id o a 4-1 TJ Ul cn c 0 c Id E-M CTTJ QJ td 01 01 E M M fd rd x: TJ a -P OJ 4J u 01 (0 a x: o ■ E to fd O 3 I O 3 I 4J 4J 1 C «P I o a) fd ■H x: M in 4J TJ tn 3 IP OJ u o x: to *J ■H TJ TJ c x: UJ u rd -H ■p x: O "P 3 M TJ "-f OJ 4J QJ >, 0 M-i UJ QJ 4J M TJ U QJ Ot TJ OJ *P C > QJ ■ H ■H TJ 4-1 TJ c OJ QJ Id fd 4J M 4-1 u 3 Ul 3 n QJ XI T) ■H x: 3 01 M 4-1 Ul o ■P TI rH Q) ■H U-l in c ffl s 03 X E rH 13 ■H u o 3 (N 0 -H Du s s: U 01 u Oi M <^ tD E S M J i-H -H in 1 4J c -u tn j:: J w •'^ a dJ tr • « O U 3 J > O I X 0 t ] 3 XI 0) M r : 0) 4J 3 x: J X W 0) 4J ) 4J -o x: 4 * C ^4-* -U 01 ( J U) (B O C ( ) M 4J O E ^ w {y ^M rt 1 ] c JJ x: w > J i-H O -H JJ O ( : fl M > : ; c w H M H ( ■rH C XI fO ■!-* t J W-J O ffl (1) (0 .r^ 4-» u-« c I < dJ -p a M : 1 x: a 0) ^^ OJ ( 1 xJ O U 3 4J ] I u 0 ■-< J ; C XI fO nj t ■H 3 C C 4 w) 3 -M u o : 4 m n-i oj I) to o ■•-' ^ -1 -p -> c u c w H ra c TJ m D -I 4J at -H u ■ -) Vj 3 4J V> - 3 o tr 3 w -H ■ 3 a , U -H U-t 4J ' W ^ -H 0 I-H rH OJ C J ffl (0 a >i-H ■ u jj +j tn .-I U -H C C TJ - H > flj at o »4-i en o ■ 1 OJ c w u H V^ 01 QJ 0) ^ oj > u e . X: 4J H -P w ■ ^ 4J c -p m -H ■■ : » M C OJ nj > W 01 M XI 3 ' QJ TJ . , .1 OJ nJ : M t3 (d o cp 0) TD 01 0) M -P u c 0) in I-H n] .-I JJ rH r-l ■H in ■-^ rtj W Ul Q> ^4 U O > M -H - y-i 14 tr • (0 V^ 0 0) TJ x: OJ .-I U i-H > O C 3 r-l QJ C OJ o ^ -p x: i-( o -H (d u E OHO c cr 1-1 0 0) H-i 1 O w E : -H -H o ti ] j:: x: u o 3 t^ 0) t-i OJ ; 0) rH -H O ^( -P 3 O m 3 QJ 01 > x: OJ 1-1 -P >p in jd 3 4J at T) 1-1 i-i c w -o a o -H 01 c 3 -K 13 (0 W (Tj >i d o w J W C O O fl 4 OJ ffl -H ^ JJ (0 JJ rH -H I to W n3 fO Q4 I 0) (d c fd ( ^1 4J U <- a >,4J n3 X M c x: QJ 0) OJ 0) 4J x: > H 4J U -H U E OJ 4J -H at j:: ji: u ^ -p -p 4J OJ OJ U) -H O ^M 13 >i 3 «-( I i/i >, OJ C W) C -H (d c fd W QJ O E n] 4J lu -H O O 4-* OJ O I o c i4 fo x: c in O Q- o c ^00 C 3 -H D Ul -P O OJ U W CP-H Qt C >4-< 1-1 fo at xr-o c -P «-( c 01 -H U O "d 01 rd -P O w -p ax: (d ip in at -P 0 01 u -P t-i u o X at OJ ■H H 01 P x: fd -p ^ (d 4-1 4-) 3 o x: M 0 4J C OJ X] 0> U) ■H U i-P u OJ u n X3 OJ 3 > 0 4J « -rH 0) O i4 t7> Q) rd 0) M >P ■P O fd 4J OJ c 3 C -H 0) -H o in ui ui ■H 0 n) -H P O E u x: 4J M U U -H fd -H 4-1 OJ 4-1 — C tn u Ti < tn M Vj QJ Id O n-i tJ'4J U 0 M c tn (d < -H rH M QJ rH fd -H OJ fO , C 4-1 (0 1-1 o in 3 QJ Vj 0) 4J a4J 0) c (d x: -H 0) 4J 4J x: tn '44 4J -P 0 u o C QJ C OJ • ■H 4J U T3 ■H >, l-l M QJ O -H 3 O l4 --I OJ 0 U rd Qj M tn 01 x: X 3 H in 01 in (0 tj OJ x: 4J tn -H vw > 0 «d (n 13 01 0) -H QJ > Id QJ 0 4J 0^ u -^ c •-* 01 'H 4J rd H TJ rd > - C 14 OJ ■ I 01 o x: • 0 C M OJ -p s ■rH c OJ fd x: — 4J C O X: TJ 4J TJ ITJ ■■H -4 E-t C C --H D> 0) 4J O C nJ CN ■H -P rd 00 fN •-H O CTi • OJ — la u -H E w 0) o . P4 x: >, 01 -H WD w u +j in in -H OJ Z 4J o TJ • ■ c --I u u -H d; tM in o OJ OJ 3 • E E U Pu ■* 4-J OJ O U W • O C E i-i-H 0OO' Ot rd O 1 Q) U4 0) 44 QJ ■' 5 U4 -o 14 4J I ; QJ 01 U O 1 01 d) ---f I OJ 01 OJ p u x: 01 j= -H 0 "J 4J 1-4 4-t TJ 0) ( u^l nJ D. (V E M Q U 15 SI TJ OJ ■H 01 c rd S CO s: E rH rd QJ •H Ot rH 0 u rH fN x: ■r4 H S £ U OJ P tTi p (d rd s za-1 I fd in C 4-J 4-t 0) U P ' ■H M rH E -H 0) ■ 3 H c 4J x: 01 tp rd CJ 4-t fd ' c ot P o - c o p - fd "d 0) O tJ -O 4J »44 o ■-4-4 P C C O rl 4-) >, rd rd < 4J 01 fd p rH jz: 4-' -H rH -O fd rt QJ rd rH ar30JEPx:x:o 0PM-HQJ-P4Ja QJ CPE-' 01 x: a QJ 4-' QJ x: 13 13 U P O CT" 4-t C 13 -HO (d -H (d 0) ,-H 4J UH QJ C 3 x: fd u x: rii ^ 01 4J P 0) 4J ■H c rd w OJ >-* x: OJ 4J -P 4J c -P c ot rd ' O 13 -H E x: QJ -p c C 4J E «JTJ OJ OJ ot tn (J-r-i P -H > -H at XI 0) 13 O P rd 3 O O E O^ 01 H4 0) a E W rH O XI OJ ai XI OJ 4J o 4J rH x: e rd M p rH 4J 0) P 0) 4J E > ffl ' 4-» 4J ■ 0) c x: H c ■ U O 4-t > OJ o 2 rt: _ CO 3 P P QJ 01 OJ -rt x: -O > 4J U4 C C O QJ U 3 01 Cr U c in c 4J M 13 O QJ O Id I E u u x: XI 4J 2 U O OJ c c fd p x: ■H QJ o o o ■H Q) >i JZ Tt P ■ 3 • O >i tP QJ 4J 4J rH C P rd ■H r4 H 0 rH C OJ 4J E 3 3 jj jj tji E OJ C OJ CTH P 4J fd fd C OJ 01 C I o rd ' 4J 4-1 fd C4JQJ^rdOJPOJ - •• U U C C CPXl fd o o 01 QJ QJ -H 4-> T) CLx: iJ ^ c ^ 4-> fd >.-H 3 _ _ C 13 O £UrHCP0T)3 ■H QJ QJ XI C rH CP W 4J (d .. rd fd 01 C >< OJ - - - - >, 01 P 4J ■H 44 rH 144 P QJ T) ^ O X] 13 -H QJ 13 QJ rH X C ■ ■ ■ ■ OJ O 4-1 rd in 3 4J C a rd CP O rH OJ 01 0) OJ QJ fd >,0CPQJPP'*hC p 4J fd -H fd U4 o fd E 4J 01 QJ H 4J c fd P P > ^ QJ0l3fdQJ-HWfd ■H -H CL j:: 4J 0) c t3 P TJ P O fd P fd rH 01 O 0) E QJ OJ 4-t 3 > MH O 4-1 x: -H o -H d uh H c p > U P H OJ XI -H 1 01 X -H 44 OJ 0 fd OJ P o E tJl rH U 3 >. 01 >i*P 0) C rH U U c -p OJ ■ -■ " ■ ■H P 13 QJ . x: H ip . 3 OJ c p 0 x: (d fd c 4-t O X Q> >P O C 01 3 OJ 0 a 01 3 3 a E 3 C Ul OJ C 01 «1 O «H C OJ 4J p Id o x: rH ^ -^ 4-130 r-i > i^ V> XI 3 C 44 QJ >, >44 QJ Id rH rH "O P H C C "AH 13 'O E O Id 0 ^ P QJ ot QJ tn tr 0 x: x: XI 4J C 3 4-1 4-1 3 ■H rH C > OJ tP P rH p x: c o -H OJ • 01 4-t -H 3 x: OJ 01 13 01 4J E OJ E fd 01 4-1 -H Tf O OJ O -H » cr> p p p in OJ QJ 144 4-1 x: 4J P c E fO u -H O Tl O XI -H E 01 4J c p rH x: -H x: 0) (d

tn -H 01 4-1 I 4-1 U C 3 OJ O X3 "w ■H 44 OJ ■P XJ ■H TJ -H OJ 13 rH C X OJ 4J • x: OJ rd •H0J0Jfdt/]4-IO4J rH C E M 3 C >,144 P (JJ 01 13 OJ > 13 ' . -- OJ O -H rH fd a > p c c c P 0) - at o < 4J > P 4J fd TJ OJ Q) #H VI -'-' ~*-' •- ^^ ^^ fc-i u/ rdoixrcipQiaocp OJ OJ E-* -H fd -H P t, V* u cpx:o-HOO0J 130 313U>>4-)HP 01 P • rd rd C 01 QJ PdQJTIOJ x:C4010lP rd ECx:-P -unir- a 0> O rd 4J C 01 C U 01 . d^33.H53d rd O Cr> P -H rd 13 fd P P fd o I QJ tn 44 > C > c a- -H QJ (d P -H 4J Z -H U fd OJ OJ c fd 4-1 QJ 3 O > tp-H rd 44 Q _ _ tn OJ 4J QJ QJ x: 13 MC-H4J0taiPi::4-i oj tj 01C34JQJ4J ottn >,0'd fdx: cx:o 4JCCLPW-P '44-H4JX: 44(d PrdtOUO P* rd rU rd OJ 01 "P 3QJQJX10lPOrH J E x: -H p o rH ) ..-HPPC4-l4-ird P tP tj OJ Id 3 IOJOJtOWE-H044 I > P M QJ 01 4J O 4J-H0J MTJPOOl 4J 4->3l3 ■HCPE4J rdTJCO0JrH0J3QJ'HP x:cojx:oird>crccro 4-Jfdin ocratu oj44 0) ^aHx:ppp44 QJW0I4JO44 3 QJ Nin OP QJQJ013 ■HQjoicaoJSx: ^ ^ 13 H 4-1 x: fd O P 4J 4J a fd fd Q rdO tnOJ4J>, POa OJPtnojx: 4Joioio)E P QjrH O 4-> tJi-H 4J -H O ■H13 CUQJW>iU QJCPfd 44'H-H0J(nrH SC4J>,OP'*4EfdrHQJ ■H QJ 4-t rd -H fd x: 4-'l3HCaU4J ^4-t4-> fd rHO0Jatra4Jd H o H H p ax: u OJ o 4-t4Jj34-) aoi4J<: E-p O QJ 4-1 13 d 01 -H -H 4J0lrHCC3X:01U -p n 3 3 o 3 x: a fd tn a a-H O rH QJ d 3 on ■rd-HOoiPOJx:fd 14-14-14-1 E — w QJ OJ 01 I > 01 P rd I E QJ *« OJ E tn

4J ■ E tn i o P 3 p ot ; <44 4-1 x: OJ H c d u 3 x: E o iH ot 3 3 . 0 - - " 01 r4 O U P 0 P - d rH H QJ *P fd 3 O O 4-1 p 13 OJ o -H 44 rd H c u cr C 4-1 4-1 3 3 -H a o -H cr o E ip tno4-iE0Jwoo < 01 I P P 01 rH ■H p c d 3 O C 34-lj: >13 >i01X) 3 OJ P " - w inPOJx:Qj>iai-H fd U 3 O P rH 4J 01 -H 4-1 3 rH OJ U EPPfdoifdx:3 4J 3 U 01 4J 4J TJ CfdUd-H-Hi300 tn OJ -P c x: d o p o ■ a o fO o 4-J 13 d 4J . Id u ■ p p (d « 4J OJ d o < QJ d E P OJ U4 M VI u ■—■■—■ 11 [JJ"--! w H a E o inEuxi rdE-'-'^dP c 4J OJ-H tdOJ -HrdAJOQJ4JOtPU UP134-1P- > -HPOJPCrd ojoto)WfdW4-ippPdJx:-H-H44 cPrHOj ot-Hoioj a-H p > 13 I c ■•-* "4-1 oi>o fddco* idotM3rdoico 44E0Jfdx: "0 ' " ■' ■' P 4J XI 01 OJ QJ C P TJ C 01 C O 3 d -H I O OJ TJ D. U d to a £ 4J OJ QJ 1 3 U P fd OJ XI fd 3 tn 0 fd C'H E o a o O fd M lJ'4J 4-1 4-> a tn QJ -p x: OJ OJ E c P 4J x: o -H (d P jc W U fd r4 rd -H 3 4-1 OJ X] a 0 tr u — c 0 4J 01 g u P OJ 3 OJ fd P > U 01 o ^ rd E CP Id ■H OJ 4J P rH CP C O fd 01 — 44 p P 01 O ot . 4-) E 4J in 3 O -H x: ■>-* 0) 3 OJ ,-1 E-t > 01 H d C d 4-1 rd ■H E U X) — 01 P CM -P 13 rd rH — H C -P fd TJ 01 o j<; fd 44 H 01 C i3 4-) fd O 13 rd 4-1 -H Id QJ > 4J -P 01 p tn a fd O QJ 4J O TJ a in H I >, O -P 4J t P TJ TJ TJ (fl I 3 O CP d d P P TJOJgCfdrdOJQJ QJ 4-1 E H TJ 4J r4 01 4J tn OJ C *P ■ H C 4-1 P 01 3 fd (d O O 01 C -H 01 p -H 4J in 0) 3 H »P QJ 4J E C E 0 TJ a 01 QJ OJ 3 o 4J x: > 0) c 01 p 4-1 O >. 3 O rH 44 O P rH O H xioaoja'H.H4J ■H P C E "3 P fO tn C 3 O -H U QJ -H d o a a H 01 4J OJ -H 4-t rH 4-t O lW4J4-l01ldHld(T' QJ U fd O U rH 01 13 at x: a-H o ^p c rH 4J tji a u >, QJ 13 0 01 QJ rHOlOJdrHrdrHX: ' --i E 10 O *P 4J rd QJ -H d QJ OJ ux:cpoJx:xiPOJ •H 4J OJ d u M 44 p O OJ 13 13 rd -H 4J 4J rH rH E 4J 4J 4-1 4J P OJ d v.ox:OrHooj: OJ a 4J H H 3 3 o .p^ CL 4J 4-) H o344ac4JPx: 0] 0) H O 3 H M 3 C ID 1-1 o 0 c tJ^X re X 01 u w c OJ in 01 re QJ 3 (d ■•-■ 4J u ■H c re C -P f-i u 4J OJ QJ QJ re > H 11 > X > 0 4-1 OJ Q> C Wl 01 U-l (fi -r-l -4 QJ p H 0 re 0) X ■H c tn re TJ 3 H 4J -H 4-J 3 OJ 3 i i O X cr 0) 3 X 4-1 a 0 C in X W 0 JJ +J •wx E M H 4J 0) QJ C «-i 0 ^ i3 tP 3 X cr4J 0 - 0 0 QJ c tP u QJ U r1 4-1 in in c *M )-■ c in vj -H re r4 OJ 0 4J QJ 01 U TJ W E 4J -4 C OJ QJ U OJ 3 OJ c (0 o « a> OJ »*J u m V> 13 c OJ 4-1 Ti ^ 0 u 01 11 re 0 ■H 1-1 M-l O r^«P X Ml 0 3 M >.E re X Tl 0) in OJ E c re ^ 4J C ot 4J QJ C c P E re Ml E X a4-i 0 H ■H .H ^3 0) 1 C 0 11 re T) 0 ■ c re OJ > tn a re 0 X OJ 0 -H P 4-t > lU (0 0 li O 3 CJ ---l a H re u 0) -P ■H Tl -4 -H QJ V4 TJ X U tn 01 OJ c E -H •H C tJ> u 01 c E-H E 3 u 141 M CP U 0 c Vj 0) 3 'w -H P TJ 4J c Vj 4J X ■H (0 in [i1 M C XI 0 H 0) ■H cr QJ H C OJ ^P re 01 u cr 0 OJ 0 01 U 3 re re re H 0 ■P t) 0 c C -P TJ Cn3 OJ 4J d H Ml a QJ 01 X 0 4-1 ■H 0 0 X c 01 u a U "H W 0) re D 0) -H Cr^ tJ re cr-p >p QJ 4-1 >, X M ^4 14 4-1 u in TJ X >i QJ QJ P Qt u H a) (0 en 4-1 U XI w t- 3 c •H at u re 11 C Tl 0> 01 3 01 in u 3 OJ c 4-1 0) »4J c c c JJ 4J o ■H UJ -H JJ 11 re ■H OJ 3 p >>A at V4 0 4-1 E M 4-) U E P 3 01 X re '^ X «-i 0 OJ 0 -H QJ ' C re M in -P 0 4-1 to 4J c 4-1 OJ i-1 re TI T) > 4J M P ■H C (y (tJ -H 01 a ■H O r 0 E c ■H re TS 01 m TJ a 11 p 2 OJ 3 a X re W M 0 x: 2 w u >t-l u-i E u c 11 ■H 4J QJ V> OJ to W -H ot 0 u X 73 3 « U X in 3 ■iJ re (u 0 0 JJ 0 QJ QJ 0 a-p OJ t o re X3 '^ re TJ Tl X c re QJ 0 10 re re. re 0 -1 O £ TJ »i -o U X ■H re QJ 3 s 3 V4 Eh a rH U C in u w ■O £ ■p tT' re C P U) OJ X u X H a OJ o in -H 4J Cl E 0 Ji QJ -H W4 M OJ tn re flj 4J W >i ■H cap OJ •rH X 4-> re ■p X a to > M p 4-1 X re o o> ■1-1 11 u 4J OJ CJ u tn 4J jr W C IM ■H (i) c O X u V r1 re 3 M re DC M Ij 4-1 • QJ X 3 a QJ 0 U 4J 3 OJ TJ 0) re OJ E Q 0) If) p OJ Jj a-P E « W X Q, OJ ll OJ p s V4 ll m 4J O T) flj 0) u e c -H E 0) ■>H Tj Ul -H c OJ aTi rH en P >i Tl a re 3 u-i tn > > o OJ jJ OJ -P ■H 3 0 QJ 0) OJ in 4-1 TJ X • X OJ XI re u C 4-1 Qj re M-l H QJ re o ^1 0) ^ 4J U D >, 0 ^i E M -H Ml t-1 4J a 0) d re 0 4J 0 c re • en c; u M 0 TJ ll X 11 aTj 0) re re cr QJ u in jJ re -P 0 re 3 3 !i 10 W to 01 to c < en TJ o a- re fl) x: -p Q, u u c W 4J M M QJ X 4J .-1 re re +J ■ o re U' c OJ i/i axa CJ in (.1 ■-< C X 0 X re -H re 0 0 X 0 3 JZ V) A-> O in H re OJ (0 OJ OJ u OJ cn 0 0 OJ c 4J 01 M-i 4-1 P -H M-. -C 4J c >i4J o -p -jj re ■p re x: 0) - >-{ > p a in 0 X U) c Pi a u c i -p 0) o u o tn X u Ml re to QJ w X (n in TJ OJ ■r4 re CJ 3 0 01 4J XI CO E U to CJ E AJ 0 P 0 4-1 a\x M H tn 4J 4J ■H X c tpx in 3 p cr 01 x: re dJ ■r^ u c M *J TJ 11 c Ml ■H CC *J 0 u Ml X 3 01 OJ E 4J ^4 5 c u u 3 -p e 3 re -d 1-1 1-1 i-l u 11 OJ in a QJ c re U < 0 in OJ OJ re 4J in X 4-1 4J 0 0 ll T^ 0 i/l -P 14-1 OJ 0) QJ rH OJ re 4J 3 E 0 ^ ■H U >iH 4-) I-) u 4J C QJ re 0,4-1 Q) u m u 0) ■-I x: > 3 c a •H o -P E ■ H TJ 4J tOTi 0JX4JXITltO OJ OJ in T) E X a -p 0) a 01 u) re E OJ M 4J 0 ^ re QJ OJ C -H C 0 4-1 M c X H re ■H M X c ll O C E 4-) 0 TJ 4J iH o cr a H X O 0 tr X Q D nH 0) U 3 QJ re 01 re 4-1 E in 01 W 4J -H ll u re 0 C 01 UJ -P l-< C OJ CT -P u (/( *w 4-t re E XI X ax in X 3 ll u QJ Qt re ^ W C tJ>H Q OJ P c U 0) 0 QJ Q) E CJ ■r4 4J 4-1 QJ in • u QJ 4-1 -H Qt TJ U 3 D ax c (u c x: 01, 01 C -H 3 X o 4-> X - 0 X M 01 ui in E -1 U re U C 0 O 01 0 4J 3 in ITJ E 0 -p . in X O -H E E CJ -P (fl re u ^^ E 0 +j V) (/} re 0 X -H X U re c -H 0 X QJ CJ OJ >iX 0 w re CJ ■H 0 1-1 •H r-^ to CT>X1 to X ■H re 4J C 4-1 ii c OJ 141 c yi 4J u ax OJ E 4J CO U 0) < -P u U >i O X p re c ■.-< M O TJ OJ 0 w 3 re U) 01 ^ 4-1 3 0 X X r- x: tn 1.) OJ -H to u-l O re ■H OJ ■H to Cd -H ^ X 11 4J 0 -P 01 X OJ OJ tr Cl U 4J C 0^ 0) -P 3 1-1 ^ IH -H ■P X > Ul X OJ 3 QJ re 4-1 4-1 c o >iX 4J > OJ C X 4J TJ .-H -H 'H E x; • OJ 0 • a 0 w C X QJ c AJ r1 O OJ 3 X >. X X Ui < E 4J re 4J in «-H 4J re tn ■-4 re c S! ^ ^ rt) +j >i p x: m u ax 0 3 u OJ 0 ax El H tfl ■rl 01 X > 0 X re 3 C -H XI U 4-J OJ -H re 3 +J ■H Ul AJ > J~» to 0 p Qt 11 C C (/) in tP (n 0 ■r4 P 4J Ui OJ r-t O X < K --I U-l ■P 0) w re c 11 c — > w E >T) OJ -H OJ re C OJ 01 4J TJ Tl M ^ M rM H CO T) .n OJ p re (0 .-I re OJ X ■H OJ X 0 (N .1 ll 10 m QJ .1 X 3 en Cl C ^ [J re ••H (/I 0 > >i C D> 3 S 0 v^ C 3 11 p > CTi-H -- 01 -P QJ QJ ot 4-1 tn 4-1 El OJ OJ c a 01 3 4J 2 x: H re u re c a c-o re Tl o y) Wl OJ 3 in X c > > X OJ 11 OJ QJ a X ll C H O 0 P u Q) in OJ ax E cu 0 OJ li u 4-1 3 aTJ ■H ■H •r4 in ll ^ re re > V4 in Ri 0) O 0 OJ -H x: re X X u 01 ■P U X X 0) 4J o U O 'M 0 X TI 3 k4 X QJ re H QJ X OJ M re S S C^ tn -P &< -p hj *J 3 -U 0) (0 +J M Z ■p 3 -P Tl < (0 re X 0 4J OJ re cr-p 3 TJ E 3 2 4-1 3 TJ X tns C -P .H X M OJ •1 -H u. S X u OJ u CP u re re S E tLl TJ 1 ■H QJ to r-f X OJ X -H C 10 ll OJ X M re 01 0 ■P TJ < 0 3 3 « ^ (X ip c cr ■H 3 U ■H QJ X -10 2 in c X 4J QJ X OJ TJ 'H 4J 3 to U-4 O QJ Cl M-l 0 C TJ P Ml M 4J 01 3 U O OJ TJ U OJ 3 M OJ X C QJ 4J cr u M-l VI 3 re ui re OJ C "P c Ml TJ in -H 0 0 TJ W) Q) C c ■H C E re 0 in u 4J re OJ u; E U H -H re M -H tn E C to X X OJ >, >i 0 QJ 0 -P X u •-* c P o U 4J re re o in u a - ECU 01 3 10 Ml ■H re 0) U 0 TJ C -r4 Vi 1 0 Wl 0 0 a Cl o O OJ 0 H Tl X ■'1 a u Ml 4J OJ « P u re u re 0 01 cr u c OJ c in X c ■H "O •-i -H 4J -H C TJ -H ■-H 4-1 > O OJ w 01 TJ U Ml -H V4 a c 3 OJ CJ 0 .H X 0 0 TJ U o ^ o re'-iTJ0cii>4J O C --I •■-{ r^ OJ '-4 >i C -H (fl U QJ TJ QJ rH E -P OJ 4J U TJ OJ P OJ rH Li --1 U Wl 3 OJ re u o X o re 4J X 3 -H 4J re w PC X >i u < 4J 4-1 ^ re C ^ ll o 14 01 re a4-' re E ■•-' QJ C U QJ Wl --1 0 OJ X in u tl a 4J OJ X 01 -H TJ ll X 4J CJ QJ 4-1 re P o c u re Wl V4 OJ X -o OJ a OJ P 4J 10 ji; QJ to _ _ re X M -H i-t . E 4J [J 4J Ml 3 4J TJ >i O C n u >, re -1 4-1 3 C -H T3 iP O Wl OJ ■H 0 H QJ in E T) X I 01 Wl ! o at X -H ■ I N 4J >, QJ a-H ll M c i OJ o _ . - ,1 CP 4-1 - . H re 01 M c re re -1 .H U U-i 3 --1 E TJ 0 X 3 - a 01 >, tn u 2 re Hi Qt -H n p 5 -H M p 1 OJ re u " X • OJ Wl p J -p e c TJ - re o QJ o re OJ 4-1 1 M -H E E C 3 M X 4J -H < 1 O O C OJ tP P 01 X OJ «P 1 -H a4-l MO D r-t 4J 4J 1 -rH M re Wl QJ CP 1 ll OJ :3 jz c )} M 0^4J E -4 -H CM 4J n (M 3 OJ X c 10 3 O X > O O OJ W) I > c : o ■ 01 OJ I < TJ Ji OJ [iH 01 re 3 E 1 X E -H 4J 1 CP Wl Cl a re Poic-i'a>-ixw)coi 01 OJ ^ CJ -P TJ a I 3 - OJ CP--< ■ >-x c u re El o c E -^-^■ I TJ a CP-H <-H t-H : tn • c >i I OJ - ni: X 3 rex re re o to £ 4J X X O M , 01 -H X ui 3 TJ C >• OJ QJ <: ~ ■ QJ Wl OJ 01 Wl U E O P 3 X 3 O 4J 0 0 W) TJ 10 QJ Wl 4J OJ re U 3 QJ re Wl E 3 5J re OJ o X ■ I E -H 4J . I O X ll I -H O -H «1 IX U QJ -H ■ ax I O 4J TJ - ; -H u 01 . i P C 3 ' p re P o X X U 4J CP 3 C Wl OJ 4-1 *J Wl u re ll QJ re Wl ■ c yi en 4-1 01 c -'H U CP-H Ji PC c re 3 CP re ,,.. _„-. .-PXCE X O -H Wl Wl -H C -H 4J330JWTJ 4-lrt:TJ-P'-' >. C C 4J ui 4-1 -H (0 n] 01 -H re 01 u n X QJ X 3 >i p Wl a C TJ 4-1 W QJ - ■H -H O Wl 01 < 4 J.; TJ TJ X OJ ll 4 c M 3 p re re 0 0 a OJ -rH X OJ u 3 0 re 0 a P cr Wl X c X Wl 3 C 4J O O OJ 3 OJ C r U 3 ' a o ■ - I u ■ T3 p re re ; re N-P a P j=; m t ■HO - 4-1 QJ 3 re i re 01 CJ U X 3 4J tP O TJ C 3 4J 3 .H p -H P 3 H P O Ji 4J Cb 3 0 *P C Wl 3 3 -H TJ ! rexU'H.-ix;-H QI0J4J-H3re3E J X U E o o 44JrereO3-H4-l0J C QJ C CP 3 X JMireooTJOW [joareoc--H TJ : c 01 re o --H J Wl re re u • 3 c aTJ TJ at w) o J O Wl nl QJ QJ 3 4-PreXP4J4JU ^ p o 0 re -H ?■ "• 4 re w 3 3 c E J C OJ Wl -H p I ; E I re X OJ p . I E -P P "1 ( I o (u I >i CPtP ^ i -H C OJ <11 ■ r-l -rH p 4-1 I' I 3 > OJ C m .H X -H ■ CJ 0 4J I P in QJ ■■ J -p re E -P J jj .-4 -H nj CP in P 3 tPX C - J (13 CP OJ 4-1 -P X < a CJ p > 4J » P tP-H P J 10 01 C ri nj I -H P X -H 01 J X OJ 4J TJ U ) 4J > C 'H OJ -H : OJ o re p X TJ QJ : ij 4J p -p u X X w: re C to p 10 4-1 : re X -P p re Ml -H I at 3 OJ 4-1 n O TJ C C < C < ■H U. - < in Wl p Ml Q) QJ o c > X OJ C Cn4J 0 3 re ■POX Wl p 3 c o 01 X P E -P O QJ TJ E U 0 QJ POP 3 Ml Wl : M o Wl 'O J 3 P OJ C -H TJ > QJ 0 > a 0 >i CJ c ■ re C CJ P Cn P OJ OJ I X c p PP4J -HH aOJ tn 3-P4J - - ■ tJ

i- OJ 'H Wl re TJ X CJ -H X 0 o ■ 4J 4J re CJ 4J X r C -H OJ 3 I X P X c I p re X >,'H ; re p re Cl I a CP X TJ p o 4J O 01 X 4-1 in 3 P O ^ O O X ' XXX re in <■■ X I P rH P I c ri 0 I QJ rO Ml X 0 OJ at ■^ a > -p : c E -p c I -H -H X P I X U Qt P 'P O I c P X a Wl TJ o CP c OJ 01 P X ■H > re QJ X X X Wl re X O X 3 E 3 O OJ CJ P X OJ H TJ 1 OJ X P p re TJ re TJ c acre QJ re a E Wl X QJ in c P W) OJ re o " u c X re X X a p to re O CP-H a QJ I E c X I c to QJ M : E W : p o QJ re 4-1 QJ Ml p -rH QJ 01 re re OJ > P X X C TJ 4J C at >iU Cl X re X X 3 QJ M P O X U 4-1 Wl CJ 3 Tl p OJ re ) OJ TJ X 01 P 1 > X X P ' ■r4 re 4J QJ J • 01 in X I 1 U TJ -H C C ■■ J CJ QJ ore' : p p X u J H re M X I J OJ a u X J J X CJ 01 re J P • M-l 1 "-I Qj re Wl Wl J rH tJ OJ P ) -^ tf ■-( 0 3 rex >ix ^ Wl o p 0) P C TJ a 01 01 X u d re tn pre 01 re re ■rl Ij 4-" 01 o P p c > cr p QJ < re QJ rex X c a c p re o o P TJ u ^ X 3 01 o P X QJ Wl in re E Wl >, OJ -P QJ X P C CP u - - 0 X o a ■H Wl OJ tn X H TJ El O C 01 re ^TJ r1 < ■-' U) Qi - W X 2 X -H a tj QJ c Wl aji ■H Wl Wl re 01 OJ P X p tn u c jj 4-1 re P p ■HO 01 X CPTJ at re OJ QJ X X C X o re at P 01 n > a p 3 re ■H u X r1 4-1 P r1 C -H TI (TJ OJ U rP OJ TJ X re o-Hwio-P3tnreo Ml nj K (f) t OP 01 o c •-1 cpx a-' O QJ Wl -H P X Wl a c > p c Wl E OJ e QJ >iX O ■H O -H O ■P 01 O EXPUJH14J aTJX o ^ 6 s O (0 CP 3 0 O 4^ ro O ■p c w M in > C w ■H 01 M 3 > 0 > U-l 0) 0 ■p ■^ -p c 0) 0 (n c o J cu jr c c nj *j JJ o nJ §• u o dj ■p 4-* c u U M ^ cn ro 03 B V4 ■H o 0 CJ> JJ >, c .-t 0) c 0. 0) c J3 01 cn V u *J J (U •H O M VI nj 3 c U) U c 4-1 > ■p 0) o 01 D H n 0 u (TJ c 4J o o M IM a. 1) ■P o u JJ G. 01 M w ^ w OJ n fO a < o Q c u a 1-1 •a 0 U-i a: a u 0) o OJ u o 0 dj x) in OJ 0) m fN c J= V4 o c ■U 4-t u 0 tn 01 a -H c o 0 c tM rj T) c 0 h en 11 OJ M -4-1 C" a 0) 0 IM -rJ X > tM Q «1 ■tJ kj 01 0) ■p ■H 0 > c o c r- c S t^ J-l tH x: > ■rt O 3 (J M -H w a o u -H J-> V4 t E x: u 0 0 0 "3 ITJ CT 0 m y-i a to u D. jJ a tT> 4J ^ u^ (U 1) 3 1-1 x: 1-1 (0 4J nj jJ JJ C U-l C < 0 -H J3 tn • O x: o C JJ C 2 rg jj rg U <0 C 01 iH > 0 14-1 fO U ■P u 0) ■U • u *t^ 0) Tl Q Q> U tn u rd cn ID OJ . 01 M C -U •H kJ Q x: - Q X] O -P nj -4J jj jJ U H 4J w - d) - Q) e c c >4-l tn cn cn c 0 ^ C -a c 0 0) >, kj ■H C O -H E -( ■H u^ x: 01 x: to -rJ JJ > CM tn ■ H o tn 4J CP C C m (0 kj fN ffj C 01 -H u ^ 3 (u VD 3 0 M U to OJ in c in 01 rn 01 o a o l4-( . o c o 01 3 o Q) O Q . (N > fN 2 < (0 a > S> .H D >, c tn o u c -IJ c Cr TJ 01 c u a -H l-l j:: 01 o tn ■w n3 U3 m 3 X r- 0 -H o 0 XI tJ H 13 OJ in tn u) tn OJ 0 QJ x: o. o H kj (0 D tn o c u in 0 o at H M-i K jJ M (U ■E 10 a TJ p <0 X (n o c 01 U-l 0) V4 g 0) CJ> -C c o CP 4-1 ■H c c c JJ a V) U1 c &> QJ (U >. c (U W > nj J3 0 k4 a) a, nj u Q. (0 a x: a ■H O CU OJ TJ O -H on: c tr ja --H E :ti -o -H c -u ~ " ■ C -H Qi fl a C ^ ^J > u (T3 a Q) C u 01 ^ a- TJ 0 0 +J M V ^ ■H 3 XI 0 0 c 0 0 0 a 0) u T) 01 X n 0) XI fd p ■p nj a 4-1 c i. at 0 < 3 u jj di 0 11 U) >> IM •a a> j= c OJ c E jj 0 01 u (0 H o -p 11 e CT c 0) x: u HI -H 4-1 TJ cr> a 0) c E TJ E m TJ 0 0) x: s v^ 0 2 4-1 0 C >. ro O C »H 01 m 01 a> O nj 4-i ■H C x: H -w 3 »w o IT) CQ x; E *J C *J -H C O OJ M < -H x; fl] d ' tr u E >, c 4J rH III C c 4-1 o m 0 « (V E n 0) -H c s X4 01 ffl (0 O X «] > s: O c ""^ V 0) o> H k^ nj s: c 0 c 0 *J c « > -a V4 c Q) rd W c ^ 0 lU o 4J 4-t 4J 0) tJi - fl 01 d u > c 4-' <*-( ■H01(0Cja4-14J= U) u] a H o o 01 « j<:4-)0V4cai> 0010104-11) >!-( aJ^njoj 4JTJ-H(d ja > w c ph 4J x: («Tt31-iC. C 01 0> > to 01 V4 Of u-l > 0 c o 01 o w C 01 *J to o ■H tT" in a E ■ I -H T) a d 3 01 c 4-1 OJ .2 5 3 to O 0) C o c u x: ID v< 01 x: cr c w O 4J c o to OJ 01 h TJ C (^ to (0 OJ to r Ci' ■H s m Ul m m 4J nj c 01 fO o s 0) o TJ > a 3 x: a 0) 4J x: 4J C <« 0) 3 4J ■rt (0 0 w ■H 0 x: TJ c XI 0 4-> 01 u Cr tn 0) 01 D C >U « 01 4J > o ■H 0 JH 5 H CP 4-1 4J fO 4-1 c M QJ 4) to 0 U) E V U C C m c V4 c QJ > c o 4J H 4-1 o 4J l4 to M 0) Ui UJ to c OJ cn a 4-1 4-t >< x: o in 01 •-t (0 nJ cn ^ ■H ■H a to c o ■0 c m U 3 Q) u 01 u-l OJ (ft >, •s u 0) .-1 (0 a 01 E >' tn C c 4J 5 o 0 O 0 ■H u 5 r^ •~i 0 0 4J 4J m % 4-) CJ p O c -H 4J (0 x: QJ 1-1 0 s 0 ■H 4-1 CP to o 4J c w to *J 4-1 as 'H m 4-1 V4 s >■ c to CT> 4-1 c 3 OJ u-l lU c 01 0 V4 4-' tn 0 J OJ E u QJ 0) CI) OJ c x: a 0 o; OJ OJ 4J 0 cn Cp s u 1-1 k^ c E c a Cr> T3 >4-l a 0 OJ ■H d 0 u 4J 4-1 D> 3 4J cn (ft X c 0 c (0 tn >, 0> ^ 4-1 to c 3 0 ■p4 5 in 0 0 4J 4-» u U u-l OJ c to C 0) 01 x: > > 0 u 0) M -a )^ ■l-l 01 TJ QJ m c 01 JJ 3 c 01 4J x: <0 01 a 0 "4 ^ 4J M 01 tn .-1 c .H CP u OJ g 0 0 to to a tn ^ u m [ft 4-> c 0 E 0> fl CP o c u to a ,-4 x: c -rt o c to 4J i4 x: to . 3 < TJ in O 3 'w t< -H H C i5: o x: 3 01 o o c ■H V- W 4-1 4-> tn cn 01 ct) 01 U X 3 3 (U C O OJ O O U -H -H i-H tn < 3 1*4 cn ^ QJ 4J o U 01 C -4 4-t QJ >, C ' x; 11 4-> p-H to 0) cn to QJ C m 01 QJ h tC 3 u 4J CT" c OJ to .^ 5 01 s 0 x: c 3 4-1 u 4-1 0 O TJ to c 0 c QJ )^ H Cr> m TJ 0 c M 0 3 n c c TD V 10 C U 'H 01 01 OJ 4-1 V > OJ ■H x: 1 QJ XJ o u x: 4-1 C u ^ OJ CO 4-1 3 u u u^ 1 tn o tn a 4J c C T) D w a p to C 4-1 u u to < 01 o x: to s ■H u-l c 4-1 u u 4-1 Ql 0 TJ 0 u o QJ CT- x; QJ ■H 4-1 <0 x: 01 c c 4J x: 4-1 3 TJ -U u -H U 0 TJ c 4-1 0 TJ to j:: u ID 0 t^ E to n tn (0 "} c o 4-1 x: ■0 OJ OJ 3 U u-l tr 0) ax: > to 0 3 tn c: c ■H X CO -H 4-1 H •H c -H U-l a> 3 4J to 0 x: = a O OJ 3 Ql c -H 4-1 4J 4-1 Q) tn 01 s: o 4-1 0 0 o 4-1 C c 01 4-1 c 01 0 OJ > 4J 01 1-1 x: u > 01 QJ 10 ITJ a 4-1 ID nj c XI 4-1 c 3 u 0 c to ■H 4J TJ x: U-, 0 u 0 3 V4 3 C 4-' z 4J a XI to ■H TJ OJ c g 3 QJ c D C TJ 4-1 4J QJ 0 fl QJ QJ 3 T) 01 3 TJ 0 > Ql 4-> 01 to X tn c u to S-i to C OJ x: to 01 U C QJ to lo a JT to c T3 01 : x: 3 0 o 0 3 x: U-, C 4-1 c 0 ■H 0 3 4J 4-1 ID 01 C 4J 4J x: u (0 u u u x: QJ 4-1 ox: tr 4J CJ -H O -H 4J < tn X O a> "i-i 3 IT) 01 OJ TJ E x: -I-. 01 H 4-1 jq T) ■ cnococT)io>. n u i-> c c 4-1 -H x; in 01 (D TJ ^4 C QJ TJ V4 QJ 01 3 U-l Q M 4-1 01 ra x: .^ E )H QJ O X3 a 4-1 0 4-1 QJ IH a 4-1 p OJ U-l c c 3 < CO u x: u M 4-1 to 4-1 c TJ n 0 w 0 01 01 x: o CTi to 0 tn 4-1 u TJ OJ C 4J u-l 0 ID a 3 > E 4-1 4-1 QJ TJ ■D 0 0 i4 OJ 01 o OJ T3 C TJ c u iD > QJ QJ Ul XI U 01 x: l-( < 14 in o C , 01 u — I 4-1 tn x: I -H .H 01 ' X 3 3 CO in 0\ 3 ' O -< tn 01 3 01 in TJ l4 O TJ - ■■ C -r-i tn O -H a TJ t0lH4-'aQJQJCI^ a x: u-l 4J 0 CO cn OJ 3H >, ■ 3 -H . to O 4J (0 o . ■H C -H . 4J in QJ TJ 4J O H E C O D> tn OJ d Ql nJ CT" D> O ■ S "■ S " x: x: ID T3 O O 4-1 i CP in 01 3 •H u o to M Ql +J a > n) -H T) u to x: d o . TJ 10 OJ 01 QJ -H TJ tn 01 o 3 M m c x: E a x: QJ 4-1 01 a o ^ a 4J a (0 tn C c C -) m 0 ■-* ja ■U o 4-1 -H n] t-i : to W E 0 0) in ■H a V > o c ^ -H 5 0) C 0) P 'H 'H 4-) 01 Ul a> , 0 g p 0) >, O ■-< 01 n CU o tn in 3 > 0) >. (0 4J 4-1 x: u C OJ I c 3 'H tn ij . 0) - Hi m c -H jr rH "TJ >, > O U I P .-H C ^ n] 'H 0) -o e OJ 13 tn E a> c 0) «] i >. 0) -O ;C 0> tn Q. H iJ E (ft H fl tn tn £ c > O "^^ o o O £ [ft ^ £■ 4J Ui u 0 O c: It) to OJ a. 3 0 >u > a 4-) 4-1 E o ■-4 3 W u *J c 0) p o nj g c 0) C en 0) 1) ti s XI 0 , C OJ P P "4 to 0|4J XI C P O n) o 01 tn U OJ P C E OJ OJ ^ W 3 -P > OJ T) fl] m c OJ 0) c ax: H XI p IT] ^ ^ U O OJ Di P ' -H x: x: c J P in 4-t -H >i ; tJ OJ 0> p P I C XJ "D OJ : J E OJ *M : Tl : -o 14 o OJ > C 0) O P P IT) ja 'p 01 m tj tJ ^ T} 0 0 3 c cr- tr 0 m 3 0 OJ x: 0 0 c OJ CTx: p . p 4J Qi 2 0) 0 Ul •a e > x: XI OJ fl3 p 0 0 x: p 0 OJ a XJ XI c 0 p (0 p M U-l ft) 0 0 tn in OJ a -H c ra tn H q 7i O rd 01 0) x: a p :» tn rH 3 > u 0 p 01 -H > 0 p C U Ch u a x: nj ■H (T CP x: -a rt x: ■'4 c: Q, c tj> 01 1-1 4.^ Ou o 1^ s 5 s OJ ^ a! o u o. a o « 4~ Ol c c t- g 5 «J c 5 c "> +J u t. « lO 4J c ■t M >*- c o o o > c +J LU ■Q > <♦- 0) o liJ >■ ID u 1. *-> ^_ c o o 4J T3 c ■M 4J E n XI a OJ o V) m W lO en ■•- > .c O *J CT> cn o o Z O o « u. ^ w o . o +J 3 -.- COO O (/) Cl — ' a> U 01 5 i -s « o o Q ei: O £ I i 6 S i S J( i5 -M in -r- OJ (J e: -o (rt g.;= ■r- O **- fO (U c O 1- «J o ■*-> ■4-> >» > c C C C 2 01 I. O O 1— Q ■f- c a. 4-> E u V (0 -o o ^ > 1— u fO I. 13 («- fO ■a ■r- +-> c 1. L^ *«- C o O • /O ^i O) C\J •*- CO OT LT) ■o r^ a> ,jD i: > •♦- Of OJ C _ c O o c ■*-» CK 4- a. o w UJ a, -^ I- «♦- E O & O 3 s TJ O J — — X >> >, e Q. i^ +J O W *-» rtJ >» - -tJ o ■.- c -o ^=>»ccoc>,c -Q O ^ OJ 3 "O (O ■■-.^"0-4J (U 0} M O i/t U»U OJiTJCTlCO^-^ >iOI-><0-<-0-*-»0 •— >- O S- c u o CM-CirtEEfOOt/i TJ o o C C Q. O •*- U O >— I — C >,W1(_) o ■■-•»- cu-— «!■»-» +j 4J -a ■.- •'■■-4-> >— -t-JOl'— C aJ>Ol-Xl C-MiOQJ E-'-J-OO^tCOO) E C Q.>*- 1- .^ .,_ 5 O :3 Q. OJ -M +J (J IT} C C +J x: E -O >—>>£:■<-,£ -M OJ -cx) CO =3 C i*- 4-> .— ■I- r— - OJ -I- O T) O CTtOCi— 4-J+J E.C ■»-■(- O lO ITJ U W t-CO-M CC7lQ>-4->(0 O o o> O ■»- -t-J i/l ■*-> *t- C ■.- -M O 3 "Dfoa>Q.a)''-jr 0-l-''OC> 4J4J(7> ClUl-C*4-CU3 O QJOQ • C O O OI 4-> ■!-> Cl-S-«+J>£l/l O lO O -f- ■•-> C (U ■■- Q. OJ ui 4-> O Ji ■i-Q+J+JUOJC +j "O ■.- i- -r-j.,- (l> ,c ■O QJ 3T3 3^-M.a Ol (O jC C O O <0 -r- +J CD to irt T) ■— E C C QJ QJ O r- ■.-'*- -i- -o ce ^ E -I- *-> o ^ c -*-> S I la 1- 3 .— O C ajcoLi_fao<_)C'i-> (/)03 L.-i->UOm O+J o>3 id+jC-— ■ »— 0>J»*-4J+J OTOE '— T3 l/ITa-TSr— qjO)t--<-co>CQjo;c 3co(o3iT)t.>acauo 1- c u 3 .- «J °1f £ "O rt) o C .Q i*- m o . 1 s- o ■•-• ^ U1 ■*-> O-Cl CO -t-^I I I? o .— o <— L. OJ --;■ T3 (O J- u - > U f- c «Q ■•-> c •— •— o := E ♦J *j e I 3 •— o c C -r- at *> <*- .— f— T- |_ ♦J «l -.- E ^ i = o — at Of Q. o -- I_ 4-) 01 4J -- O ^- Q. »— g ? *J V o. •— > 1- *J E ;= D .— -W 5 5 , lU 1 ■»-» 4J v» t- •M U 3 c 9i 3 >> o 1— D- C O O o '■" o a> o c •— »~ > c 4» O O 01 U 1- «I EC (rt ^ T3 ^ O T3 ^ U *-> *-> C T- ■-; lO If) O ^ e 1- 4) Q. -^ 1- O J= «» u « C ^- ID U « 4J L. O V 4-> *J Q} CTl trt £ C7> t. fd fc. 0/ o O > — a.- o « •O O (O -^ 3 £ s ^ S 2 ? .— .,- O 4J -tJ X I- "C t- z < OJ •«- I - T- HJ -^ 4-* <-■ Of iO »rt c > >» -^ c •— -^ ♦J 41 *-» T- m ^- .— 1— c •— "O I - •— ex = ^ -8 1 •a cH 01 c VI o 5 ■4-» to •a c O «t 4.) ^ n o E c •1 o ^ c « v £ f «> *-» c f o ♦> u > i J2 *-» c 1. o -J H- £ Jl •r- t- U - ^ C := ■S3 -r- *J Ol V :t; ^ ♦J -- -.- E Q. -^ --- 0> *J « -^ r— ■^ ■— <— «J £ i t B £ I . I •— O '- ^ a. t- »o •— -^ o 1- f- o •O -t-) 4-> ^ E 5 « « C C ^J ••- o c le ■^ -^ E 3 4J ■.- i— U 4-> ->>■*-> ai ti •r- f— 0) OJ oj oj Q. ■.- +-> O. ra 4-> I— O) QJ -C .— (/» fO QJ

  • .— m r~ 4^ ,— ,— Ol CT U E *J x: ^ QJ U -^ -.- r— J3 -c en 3 c.— * i» -r- QJ OJ >- QJ .— I— -.- u ■ — 1 — C7> t/l r— C71 ■O -F- .— OJ -r- .— QJ l/l ■.- S S V 5 « 5 (3 |S i I z s Q c en uj -D i -H •f- en *J cl •-* at V oj n) ■■- en D. •<- Is? ■r- Oj O -M -— - JZ U OJ .— 4-) *J 1. Ol I — ■*-* 1X3 • — 01 1) 4_ E .^ +j +j O «— -W ^-J T3 -.- tU QJ 3 CX > x: -c E ■M ■(-> en — ^o I— E — o c J- -1- ■I- 1/1 -r- (J ^ .— •.- o QJ *J r— •— TD 4- 1_ -r- ■— (U > » «/l Q. un uj -1— 3 ^ -o (U CT> t— - — ■ OJ U- (O ■*-» o ftJ ■ Lf> ^ g -5 o ci: 01 .— •.- *- £ ^ !-• J=l jD .— i— O »— 3 i- ..- O O -ID 3 u ■P O S (D g § +3 o c E h a •3 vi a O O. 01 O (0 -C ■H n 4^ rH © E J3 O 3 C :: h -H -H a 3 h o o c «i-i W -P O O 0) tfl "H f^ t. -P ^ Q 03 o Q) e -p ■P n) nJ T3 ca -P t< H gj -P T3 p, a C 0 u o O M D, "5 I 5 n) -P .3 4) 0) J>» > Ph > f-t ra -rj +S 5ii §• P 0! S8 O -H S > §^ O 2 ^*- CD 11 ro HK l§ 5^ 2 g lULU P O (m 1 ^ 10 0) Vh ^ n -P -i-J e 5. u 1" =.fsj: 5| -P o CO ll an o •s . 10 -p P j:: ^s a Id o o £| to 5 ^^ -o ® :i 1^ •H •»P QJ $ QJ ^ ^ .^-0 E U c CTl.— -^ c TO O <1> QJ 0 -M QJ 'i QJ QJ 3 TO 4J -0 TO D. •r- C i- QJ i- in SI 0 QJ 0 OJ C E 1- 0 •— QJ QJ TO VI O O.^ .0 CM .C !o i^ 0 ■*-» 0 ■.- J- c j= 0 ^ .= 0 *-> V» Q «1 •.- CU 3 3 >i I— TO T3 E ■M-D Ji£ *J-.- QJ VI ■*-' x: CL C C -r- +J S- CL CL.— >, C O) QJ +J QJ at OJ -cz 0 0 0 QJ OJ E "O ■a s- 0 0 ■0 I- TO !- x: 1- 4- t*- Q. ■»->>. 1= -0 ■■- ,C 1 0 0 I- TO 1/) 3 O-C 0 0 E XI a? 3 TO ■*-> 4-> 0 ro "to OJ ^ rO 13 QJ TO n OJ VI QJ 3 TO **- JZ - <_> Q-t-J £ i_ 4-> 0) (J c QJ JD ^ >,VI 0 -M "^ OJ E OJ a. • , -Q t. TO c -o >, TO U 4J VI VI 0 ■<- Q. TO CT> S- -O ■■- E 3 3 1- C CL E ^ ra S- 1-1. . CT >, C IJ TO -r- TO T3 to ■— -0 U VI TO TO cn T3 V> 1_ -.- c E 3 13 U 0) 0 C 3 01 J- i- -0 QJ 0 OJ Q) ^ -l-J J- ttJ i- C7i*- 0) E I- 0 -c Q. 0 c: -C £ > Qt QJ >,--- -tJ C '3 l-TJ c u ■*-> ■M > TO >»■■- 0 Q- U i- cn TO 3-»-' OOU-^ C QJ QJ c 0 •■- -.- TO 0 I- 4- *J t- CL C CJ en l/> C -Q 3 **- QJ 4- J= cn QJ OJ ■M S_ QJ c 0» TO QJ CL OJ 1/1 TO *-> 0 •r-M- C C 1_ Q. VI 0 cue -TJ ■>- OJ QJ QJ C -.- 3 C VI c c a- 3 OJ LiJ Vl 1_ 0 to J C > JD > 1- >» 0 cr Q) OJ cn u *-> a 0 >» Z ..- (♦_ ••- f— ^ 01 n QJ S- Qj s- QJ QJ .C -r- C 01 QJ 1- 1- c OJ U 0 -Q +j a. 0 1. l/> Q) > TO - E T5 (J -.- 0 >- -M Q.+J -p "C QJ CD 0 QJ QJ C ixi •<- > - Q. C Ul "O S- C -.- TO V) ^ C VI C V -C Xi 0 C QJ -CI EI 0 QJ QJ 0 CJ> 0 QJ 0 QJ >^ C QJ QJ +J l_ ^ 3 1^ 0 W •>- -0 TO ■^ QJ J= 0 — TO 0 • L TO 0 V) J- ■ > .— ■*-> QJ (O 3 •'- >,-t-> C -Q ♦J L +J i/l QJ 1- ■— c cn 4- 1— TO +J +J C 0 */> -u ro Wl i- 1- C V) OJ -M > QJ QJ - 0 -o c 0 ■.- cn c ■■- i/> T3 ■-- O) trt'9 0 .,_ QJ +J ■- VI 4-> -a +J U 3 QJ C 3 -W 3 *-> C ..- Q) -M J- C 0 3 +j (4_ p ■ 3 +-> QJ C 01 01 •— Q) (U c o> W) -— 01 Irt fO •0 -Q ^ U QJ TO i- E O- C xi u- 0 JC Q) ■t- T3 .c -<- c ■^ QJ ■.- CL 3 +J > U 0) c I- -0 >*- -^ VIE 01 ■■0 >, C < 0 X 0 TO S- 4J a 0 Ol 0 *J .— C 1- C 01 Ol i/l J- -C ■M 3 QJ +J 1- C 0 QJ ly) > .C *J 0 E 0) (D TJ 0 0 ■.- 0 1- C S_ c VI 0 > TO TO OJ -^ > '*- E a. i^ •*- 1- 0) -C QJ 0 -M 0 Q.-0 C C -C = +J >. y- C_3 ■- QJ T3 TO +J -C cn-^ -C S- -tJ M- .— ■■- 0 0 X3 U ■M 0 C 3 OJ (J c QJ +J E w c >> • +J TO TO 0 0 E 3 0 o> *-> TO VI 0 — • > .- QJ 3 3 TO ■•- -r- W l/l ^ TD ■i- > C .,- 0 -^ TO -C .— ■0 3 E ■M CJt W Q. CJl 0 TO S- •— Ot S S- CL C J- x: 3 -M ■J 1— 3 QJ 0 1- S_ 5- tn c c TO ■>- TO VI S- OJ X 0 QJ v> Q-**- VI S- ■M 3 ■•- m c (O >»■>- ■<- -■- -C .— QJ QJ 3 V> QJ 0 ■*-> TO 0. S- E cr O) 4-J l*- .— c v»-o U TO ■ 0 QJ OJ 0 c ■*-> c lo u ••- (jO O) E -O QJ OJ i- QJ TO >, a. ■.- C C T3 VI (_) TO 3 I- CL C - .— Q. ^A Ol QJ J= cn TD .— ^A "4- 0 0 tJ ■M c c (O -Q *-> UJ^ a. OJ VI 5 OJ --0 CL i. - C -r- TO 0 -C 0 OJ 3 -rj a Jli kj Q) 0 TO vi 0 0 OJ TO 0) QJ C JD TO +J >, OJ QJ ■ -C 00 Qj irt "O QJ 3 ■>- -C *-> CLZ I- S- X3 0 3 TO.— >*- -*-> c 1- OJ 4-> ■ .C QJ > TO Q.*-) *J QJ >. ■^ 0 QJ QJ Ol 3 J_ -M QJ 0 OJ TO ■'- QJ CL 3 =3 *-> 4- +J .— U -C QJ -C -c E VI 3 +J 0 -.- 4J 0 Z XI a. VI C 0 .- 0 i/i s s- QJ ■*-> .C 4_ o. ■ -M 3 — -^ C U T3 «+- E ra c 0 TO i- 0 m QJ > > CO 0 TO > 1— -O 0 OJ 0 o> TO 0 C ■1- -r- 0 QJ >— Ol -o 3 Q. -c i_ i_ C C 3 .— ■*-> QJ c QJ E Z QJ oo c +-> 0 VI »— T) "- JH .— -C -0 CL 3 4-' 0 QJ -- -r- 0 ^ C J.C — 0 1- —. 0 ^ 0 TO -M 3 M- .a Irt TO i- V> OJ -C *J -C 0 4-> c 0-0 3 i- 0 -o 0 *♦- .a 0 CL > +J 1- u TO-C TO OJ ■M 3 ■>- 0 -C U TO VI 3 QJ c TO -i- ♦JO c .0 TO QJ U +-> QJ ■— T3 C OJ TO VI C 4-> *-> C U> J= 0 Qj 01 TO C J= U1 "-^ x: 3 QJ J^ -C +J CO 1- TO -O QJ "O QJ 0 J= QJ .0 *-> "4- I- Q.U- 1— ,- QJ 0 0 3 U J- 3 C - O- TO 0 OJ TO ■t-> c w > - - QJ 3 TO e OO 1- S- 0.-0 ^ OJ 3 ^ f— cn ■*J ■•-> (o 0 O) .- CTl CTl-^ ■*-> QJ > 3 . « QJ >i TO VI QJ QJ TO QJ c v> 1- E > ■•-> C C (J 3 CO TO i- ^ (jj J= s- 4-> 4-> LD QJ VI x: >A ^ -0 -^ 3 C U TO TO ■>- ■>- ■<- "*- 0 0 Wl C U -c 0 cn x: QJ o» 0 U Q. QJ 0 C u E H~ QJ > > C Ol QJ 4- QJ QJ QJ QJ i+- CL 0 0 ■*-> 1_ TO C TO S-'r- C U OJ OJ 3 f C O- .C i. "4- U C TO QJ cn-c TO JZ QJ S- cm- -i- W) E E V 1— ■■- 0 l/l +J TO O- 0 TO a a. Ol U 3 TO ■•-> C +-> TO E -O 1- CVJ 3 t/> ui ■— •— ^ CL L. X3 O QJ TO ' — I C^2 VI TO C5 2s- ■"■ , 0 ■4— ' i^ • TO CM c^l 2^ QJ Q> Ol CM T'g ■<-J r. ^ ^ VI 0 C S- on (_J TO -i- r./ E > ^ cj >^ 5 c 0 ■^ * E LU 0 -*-> c ■.- >*- C\J c 0 .— 0 OD r- E cn +3 c ^ QJ 3 U E ¥£' 0 TO J= m- • «4- 0 0. V) Wf 1_ <*- ££ OJ TO ^^ e: 0 - — ■ Q 3 c 0 c: *-> Qj OJ TO U > 1- S- 3 QJ 0 V> VI -»-> C OJ on 0 q: 0 QJ i- c 0 -f- g-'- hi E Ot E E ■i- cr TO cn c QJ -r- C QC > 1 OJ IJ 0 S u t- VI -tJ > VI u c 0 I- LU CL TO TO C >*- et S- J_ 0 M- 0 **- 0 r— -a I'l C^- 0 QJ U C 3 O) 3 i- CL x: C OJ 0 TO 0) ■>- TO +J -C QJ ■■- U C 0 TO 4J 3 > C ■*-» T) TD 1- ■*J 0 T3 >> VI QJ -C 4-> QJ C C S- T3 -- C C C 3 C XJ QJ QJ OJ TO OJ OJ 4J TO TO QJ c -0 ■<- TO - • OJ Oj ^ E -M JD C C V) c 3 TO C > Q. 0 4-> i_ VI -0 E OJ >, V> 0 "O CL TO 0 QJ C Qj -Q .— 0 •■- TO TO V> W<- TO TO QJ OJ C 0 VI s- QJ !- i- (.J -.- -U 4J 1- in M- OJ -^ TO TO > VI OJ •r- +-> t. C TO C VI fo i+_ cr>4-) XI CL 1- TO -O i~ > .^ O; ^ QJ l/l QJ Z ■■- VI QJ O- Qj OJ x: s- >*- C .C 0 VI TO E QJ JC QJ TO 0 OJ 0 0 LU VI -M S- 0 u +J QJ XI OJ TO TO 4- CL 0 "O QJ — TO -C -.— OJ T3 C OJ 3 ■*-> 3 I- t. t/1 (J QJ 4J -C 3 4J (J TO TJ j; OJ -C CL »4- O) 3 — QJ TO 4-J 3 cr ■.- u^ IT) E *J c TO 1- 01 Q- LU r— -D C X O) QJ E -■- 3 T3 C QJ 4-> 0 4-> s- •• 0 S- Q .— C 3 OJ QJ LU S- 3 C3 OJ QJ 1- 3 •— ' 4-* ■'-' TO ■.- -o QJ x: C=l TO 0 0) U - E QJ QJ**- S- 0 C E TO S- VI Q.I— QJ C OJ QJ .C -C 0 OJ -.- I- LJ 0 s QJ •— TO j; QJ U ■♦-' 4-> 1- • <4- ^ QJ TO QJ i- .C Q TO 3 00 4J >l4J -C 0 cr ^^l^^"^"^ -4_ ^ aj ^ OJ >,4J 0 QJ 0 QJ +J > Q.-I- C cn s- i*- VI X) <— c .C 4-> ■0 TO ■>- 3 c >, >, OJ 0 0 3 QJ -O TO 0 4-> TO x: xa cJi t/i ■.- 4J u 3 cn 0 .— cn c u OJ >> '*-' E c .c 0) 4J ■.- TO >, 01 -.- ■— S- TO cn J^ TO TO ■>— u ■■- QJ C 3 QJ CL c > -^ TO VI s- c: - ^ •5 ^ "O -^ T- >♦- QJ 3 t- 3 0 Xi 3 -— ifl c Q- 3 QJ *J TO J= V> E +-»-Q u QJ 0 OJ 0 E TO ■D ♦J c 4J 3 ■■- i_ QJ ■- 0 sz QJ V> 3 ■<- T3 3 0 QJ ■*-' TO 0 u- ro 4-> VI C ■.- VI VI TO *-> C -C C 1- 3 **- TO I- CL ^0 .- 0 VI ■.- -C TO U OJ 0 0 VI 1- Q.-*J -C ,— 4-> -.- U 3 Hi 4J 3 C aj.cc aj 0 c >i CL 0 00 TO ■'- QJ .c XI C VI 4J X3 OJ C .— OJ -o 3 1~ ■.- VI QJ -<- C Q) E TO TO TO -M ■*-* 3 OJ LU -C - U 0 OJ C T3 C C , c -0 ■•- QJ " c 0-^ ' tj TO J- 1- -O c •■- VI 3 (J 0 3 3 TO "- -— >) 3 TO X) QJ TO QJ JC 0 0 ->- 3 VI 0 ♦J *-> . — Ol cr Q- OJ > > x: 3 CL v>.— 0 t- VI VI J= E TO 3 OJ £ QJ OJ U- -^ ■0 4-" s» QJ 0 QJ 0 QJ VI 0 ■•- 0 -M 4-1 -0 Q QJ TO VI 3 1- CL 4-1 (J t. *-> TO TO >i 0 -■— CL -0 4-> 0 V> *j i*_ 0 QJ C t- QJ .— QJ C OJ ■<- OJ E OJ TO VI S- '•- C71.C 3 0 QJ -C C 1- c E TO VI c C E yi aj -t-* +-» **- > 4-> 0 0 .^ ^ .r- -r- Qj v> 0 • 1. c i- ■;- 0) cn QJ -c s- 4J QJ C QJ ■— >1 -*-> OJ " 0 s- cn >-,-o > TO OJ -0 4J TO ' 1- QJ -C s- c >> OJ t«- TO QJ <4- OJ XI QJ TO TO 3 1- E cn Q) 4-> TO 0 0 vm- s- E = *-> 3 1- ■— ^1 Q. 0 > -C 4-J 1» C fO 4-J "O ■>- *-> O) c 0 cn 0 c 0 QJ 4J TO >. c 3 VI OJ cn Q. ^ S- 0 C X: TO U- Cr- o» > TO &- 0 XI Ol - QJ 1^ 4J TO -t-l 4J '^ QJ TO 4-J C S- ■ >!.— x: TO cn i- > x: QJ QJ OJ CL QJ C TO l- i- 00 • -.- u XI VI 1- QJ 9 -C OJ C_} 01 TO ■<- >^ cn >, cn 0 4-> Q. OJ .— -r- QJ QJ TO 1- c^ O) 4-) E -C QJ ■— i- c Q. c ■0 E E -c T3 -C X3 CL 0) QJ 4.1 ^- 0 TO ■»- 0 ■>- s- QJ QJ •■-■■- u 3 QJ 4-j 4J x; u 0. 3 4-» U 4J OJ S- a. 1 cn-r- 3 QJ 3 VI t/1 TO 1+- C (_ QJ OJ x: TO n3 ^ OJ ■*-> C 0 4- 3 4J *J 0 3 TO -O QJ VI QJ 4-' Q.-C r- i^ OJ 0 3 0 33 C OJ E " E 0 QJ VI TO 1- OJ "O C VI VI u. c c 1- 4-> TO 1- QJ QJ 4J C U- E r- +J " a-r- •a: u 0 OJ TO 0. cn OJ TO 4J QJ 3 .:»: CM C TO 0 c "■»- ^ ,C C J= £ C Q. QJ .— &. TO cn VI +-> QJ .— 3 ♦J •r- 4-> 4-> et TO E CL+J OJ a. VI TO 4J xa -o > Q. > E 0 3 -c TO 3 3 '^ -.- <4- 4- 4- 0 0 0 OJ 0 ■— 4-» U t~ .— CL«a: 0 ^*- cn 0 0 0 TO *-' OJ 4-' E cn >■ LU O CO -J < U a o _j o o N i^ Qi o K & q < to C i Ul C M i 0) -P ■H J3 -P U *rH tr* c o 0) ij (B O 01 Q> -^ TJ e c o 0 (0 OJ l-i O E OJ -l -H 0 -H 0 O C CO U-i . > >t i3 (d owe > c OJ 01 3 E o 0) 4J la •rH (tJ ■P 3 - 0) o tn ,^ U ---1 - O -H '4-1 +J y-i Ot "^-i U O (CJ OJ £ U-f C -H ■p o> o tn cj 01 U 01 QJ W 0) D> •wo O U 0 Xi 0} m o M 0) >i-Hx;xi QJ E 3 T3 01 -P fd +J nj C en c ■H >i W X >i -tc oc E -P 0) 1^ ,H j_i 3 E >i(D-H»4-i«4-i OJ nj 3El^ ojjmuoox:co U QJ -H 4J O tfl 0) ■H(d<*-lO)>i3-HO)'W -hVj OJD-PO-Pffi---* nj fd to xl CT-H ui rd ,H 1-. > -P cnm+Jutui-iig^ to OJ -H -H 0) ji: QJ ti -H cnjr i-i > jj 4-> 3 2 H 3 C 4-< M 03 TJ Tl O O 2 ^ to o o tn -^o-H OJ >iOJj:: 0)'-Hj::xlb EC:(3C'TJ4-)TJ n3 cOfTjj-io-P ox: 0) (d uxi+j Q.4JU-J E+J ojx: QJ i^CWfdO -Hi^^ U+JDOOI )-i33 OtnjJUMOJCOJ 4JT3 ow x:o-PC(dc CE-HOrH4JAJ30Hr3 M X) x; to ij\ Q,.H OU-) ■jJCCCE-P'J-iOJ XIOOJ 0-H-HOtdO" ■P TJ O H Tl U M 3-P-H-PaJ>iTI OC QfdtO U--tOI(dX10 CO x: -P jj 01 - -~ O 3 U Xi CO > 0 CP rd QJ T) en -H a C ^ >i QJ 0 -P -H 3 (d C Cu i^ >1 > O rH M 0 +j -H -H a 0) fd a (d rH jj o a 0) 1-1 w c u 3 M "4-1 Id fd 0 o MHO ax: u U-, ■P 3 C Vh QJ O QJ H -, QJ C"rH jJ o C -H fO M ■H O > a. E TJ j-i a -PAiO+J-HXl -liUVH -H4-ifdC "-aO QJOi 4-i.-( 0J3TJ -HCWC O TJ ^ 0) • 0) -H C -H -C-PUWUl> OJ-i ■PEO -HWDOJEUid C Q) Cutn -P 3 TJ 0) s QJ+J(nMWU>i JJOt 3 E tn qjijjth Mxiw tnx:— s ■'>iMQrH.H -H>,4J2 - « Q-'O to m u tn QJOCOJE MOOMDfd )-io fdx:-Hmtqx:uoM oi ■PQJ UH tO-HQ 3Wit-i — to ■P 0) -p . Tt tn « c 01 (d 1-1 O QJ 01 u u U OJ o c O 0) QJ u c x: ■H 4J O ^ CO rd >4-i -I E O (d tn E -P QJ QJ 0) 0 ji: tn 01 U -P M -p OJ tn Ui to to U OJ 3 >i-P Q) c x: 0 to c W 0 5 01 ^1 td 4J CO 0) XI 01 u OJ x: M -P !-. 0) 01 x: 3 3 CJ x: H -P 01 C 0) 4J to m «w O 1-1 ■■ o (d M n3 to ■ E TJ -P a O rH C O (d -r-i O l^ 4-1 rH 3 rH fa 3 0) c o cr (d s x: (d o w iH >i ■p -p u 01 fd 3 to 01 > . E o -H 0) x: 01 >, W TJ M 4J I QJ C td -P OJ Ti c tn u x: 0 tn uj -H 3 M to QJ O U t4-i U «4H n3 QJ cd jj x: ■P C -P tn < E >i QJ O ■p x: h 0) 4J ^H UtnTJOEQJ-POJ-HDi QjM(daj«_ictoxi> I V^ O rH ^ IH 3 OJ 3 3 - Q. > to O +J ■■-i O rH (TJ 3 fd (d Q> -H tp -p 0) CO -H x: M H w x: OJ o x: x: ■P 4J TJ J= . _ > ' C U fO v^ to (d QJ to -H <-H (d - ji: ^ j-i jj c a,M I i iH a-P u >i jh QJ jj a fO (0 O X: 3 3 MH ^•-1 E &H -P c to o ■ -M Q. > t/1 C QJ QJ O O **- >i(J c , Q. O O E ■<- 3 3 u^ O C O J= -M trt cr t. -M QJ i- 3 c Q) QJ en Q. (U 4- t. Q. C iTJ w E O ns ••- OJ W Q. ra Q. fO 1*- ^ QJ O ■*-• U C71 O ■*-> o • — C -.- C ro O OJ QJ 4-> O E i_> ■'- QJ O 1- -o .— fO ■•- c z 3 >. QJ *-> ui O 1- Q) CO 4-i *J Q> o x: ■M «^ Q. i_ roc a CO 4-> C Q) QJ TJ o o "^ tn-o *J 4-" 4-> QJ C QJ QJ -O ■-- i- 5- -.- U 1. n3 O cr^ QJ ■'- +J (O Q) i/l C X) .^ (D . — S- i- Qj -r- ro ■*->*-> O Irt Qj -r- O 3 (/) > ■■- (/I O) (_> -r-), t- X) c .— 1. c ro O 3 ■.- Ol rtj **_ OJ QJ > c: -r- cn^ 2 ■r- u QJ ro ■!-> ro -O E E QJ XI ■ c t. -,-+-> wl Qj (/I OO TJ o 1— TJ T3 3 > QJ 4- E r- O QJ > 3 -r- -1- -^ S 9 r- O I- r— 4J a. L. ■ — x: ro OJ ro O a ro trt > X) +J U QJ i- ic m s: s: Q. L B Vh 0 ■p +> 0) to jz: o to Pi h -c: cu 4^ rH t> CO S O rH o c: c fn • fi CO Q) to p> tu P Pi • 3 O P P r-\ to o to cO > >->-P Pt Pi o bO CO 4-> -H o QJ 3 Q) -P Pi cu o H 3 C CO > -d p; ft o C3 C pi o +^ OJ S P! nJ H > P C o o O O P: QJ p: a H H C o -P P' bO >iCO -P M H to H rt a ^t o CO QJ > C O O cu Q) ■d CO QJ -p x> (u p: CO P o C to Pi p; H o +^ ^^ rH CO 0) H e H QJ QJ O CI- S ft Pi CO p; H o o QJ tu d cu O 3 tH rt ciJ CU bO CO 5 w a Pi j:^ t3 to OJ H-> "H to O H CO Vl CO d Pi x; o o Qj OJ CO QJ to -H .e t> ■H CO 3 CO 3 ? -P H X> O QJ QJ H P Vl H P o s B -^ [0 3 >:; » cu QJ cu nJ Vl CJ o si o E^ Q) tH >•, o x^ Pi Pt XI H +^ Vl cu (U (u a ho O CO r-i >>Xi =H si T3 ^ U -^ r~i tu -a -P Pi Pi o +^ H ft •'H-' Vl to Vl CO CO 0) o s c: CO • QJ CO -Q rH +^ P" (0 rH CO 3 ft to QJ to 3 to P CO Pi QJ -H (0 C m tH M nJ H 'H QJ C: H C\] H CO 3 0 o bO M QJ C j:; -C! P to QJ p: C d 3 o Pi 3 Oj o Pi P- TJ O V( P to C P* V CO (0 ft CO ■o ^^ C OJ 3 -Q S -H 4-* S • cu Pi J=: (0 -P CO C! O QJ P> 0) Pi QJ -H x: cO CO P! H rH tti o > o rH O Pi rH ni +^ CO OJ s :s o cu P< T3 0) HQjQjp>)'0 cS^ bO § ■H x: H-> CO P- o P> -c3 O -H > CO ft.C H m ^ 0) tn 3 0) -C o i-H Pi -H bOH P O rH P QJ p p: p: H +j QJ p: -H Vl a ft H Pi x: (0 H rH bO (1) j:5 4^ C rH +J O 13 >>o si ■H GJ-P P! U -H >) to bO d o to o QJ +* x: P= 13 CD * >- en X a £} QJ Pi ft hO ei+^ fO cu -H O rH QJ 0) QJ Pi cO O • -H CO oJ cc; CO CO Q) O C^J to CO CO OJ p: • c Pi x; hO fH ftPOPl4JQJftJZlPlP|t03QJH Q) 0) j:: QJ P! ^ vi > si-A >, QJ O P- CO >^ o O Pi (0 Pi O O rH > 3 m X) Q) o " c QJ ttO-H QJ QJ O O rH • > ■1-5 a -H 'H o 3 c +J 3 QJ H O QJ ■J Vl -H bD CO bO ^1 H ^ CO OJ 0) -C ^ >; -H Pi CO CO O ^ cu p; CO H QJ O -C Pi Vl p, p: (u c: (0 p: x: QJ ■H H -a 3 0) o ft 0) CO tsi o a COXJ-H boojx: COP> ft QJ O CO P" tu 'H -H Pi t-H d nJ H O H H O rH +J >< CO c: »-i H o CO C p: +* 1) O Xt Vl CO rH CO 1—1 ^ C CO CO avt H-* 3 CO 3 t3 cu -H ^ o QJ Pl030^QJ0-P ft QJ Pi QJ h x5 Pi 3 T3 o H PM jz: CO ^ CO +J 'H to CO ft-H P 3 ^ QJ H P* bO-H Vi CO > ^1 v O H Pi QJ CO r-H 1— t CM fH 01 QJ ^H QJ a 4^ H to 3 c a ■p -p O P> P" rH -d p» o p: QJ QJ c p: v pi QJ x: o 0) nJ in O (IJ c; 0) 3 OJ > Vi^ Pi CO P p: CO O QJ CO to CO H to (0 O QJ CJ ■H p: > tu H Pi to QJ ■H +> O ■H fH O C H o 3 O 1— 1 to O CJ o t> rH cu 5 QJ P rH ^1 to cu tJ CO > QJ CO H X3 vi a C\J QJ U ^ xi >j-H P o ftfiq C O QJ O Pi Pi fH QJ 3 cOh >i QJ c: ■H -H x: VI CO 0) Q> j=: o a +^ r^ CO -P 3 ftQ O Xl O 4-» cu cu QJ 0) B QJ O -P rH > P> 4J P> QJ to C B -H t3 4-> VI s bO-O ■P H 3 O 3 OJ CO j:^ QJ Pi +J 'H S p: q w rH CO (U CO 0) o 3 x: xJ n5 p! til ^ c C rt C Vi >, CO cu O CO c CO ^ > o Pi to C o Pi S o +» o cu 0) CO to 0) ■H cu cu tu OJ si P> >T,M cu -y o ■P X: -H to x: -H CO S QJ QJ+J 3 O fHCO •H S M c x: ■a xi oi a s: >,+j hH f^ j:: -H CJ PI to cu bO+J tH P o CO x; x: box: CO to o • -H M Hpq ■H +^ 3 QJ QJ -P H +J Q +J. H M VI QJ a p: p; o 01 QJ --P> c; 5 J^ Q) X > VH P W > rH ■H 0) o si (X c ■P O • Ji: cu p: t3 ■H (0 CO In H C rH CO f^ G rH -x: C >> 0) QJ Vh 1-^ o P! +J QJ OJ O Pi ^ -P xi QJ H bO d QJ XI o o ■H QJ cu (0 H 0) CO O V rH cu +* c ;a nJ c: -H fH X^ -H CO C O P> (4 3 p: c O .C QJ QJ •H H 1 x; Pi Pi H -H o rH O ■H V) c o o V ^ O -H +^ -p -p QJ o Pi P> P» O ■H o -HP* 3 x: p: VI pj cucoB^oviPcOrOPi iH O QJ CM -P Id ■H CO • nj Pi H x: -H CO 01 bO CO ? 4J ■H CO P* tu Vl CO XI -H cu -_^ E CO Vl CJ 3 iH B CO hO >3 +J CO s: 3 4-> CO to E QJ p CO P" ■H QJ "^ -C 3 P Vl P- Pi p, -P HO -M CU ■H r- C cq 0) CJ QJ +J O 3 O l-H CO 3 --H 0) hH ■H c > > Pi P P to H >> CO rH tu O -H (U Pi CO S tJ u ■H jz: ^1 • o O P, ■HO cr CO XJ -p Q} ■H -H QJ CO c o Pi cOPl^iCOXlP-OXlOOJ ■H n] S ,c M +J n) >, tH t^ -p jz: S Cvj cu ^H HJ o !> -o Q) a to H rH o S O CO a P" H rH V P» u • ^ P. o w nJ 4-> p. 3 rt CT) -P p; (0 O Vi cu CO x! bD C o Pi 3 -H ft M ■r^ O H Pi c: ft CO 1 Fh 'm (U o rt 0 c c o O cu -C -H ,C (U o o Q) o p; P> Ph -H S X^Vl « Pi HJ CO u p; 3 o H ^ H Vl c^ &I B QJ ^ ^ O £:oP cr;SE n o < a to C -P S P- O O M Pi o P c CJ en O QJ QJ ■H S QJ x: Q) c pl £ Vl QJ in fO • CO +J x: rH C u z j:: c p) 4J rH 0 • u ■P Vl 0 en •H arH jn D 0 -H Vl 3 3 to rH U M 13 AJ 0 E QJ cu Pi c tn -H (U Pi 3 fO QJ fO QJ tn c CO 3 QJ x: ■H 0 0 CO CO to JJ QJ U C1.-H •±\ -- fO at w c 4J a CO C PI E H QJ . "0 >1 aj (0 tn 0 -P -H CO > D^ QJ tn QJ Pi to Pi U 0 • Pi 0 J= MOO - 14-. QJ -H 3 QJ \ ^ c ir> QJ O -P n u c 0 Pi avi X QJ QJ tn c ^ ^ ^° CT' C (0 XI 0 QJ Ti x: 0 ■VJ* CP^^ V O Pi Vl CP ■P u ^-^ C 10 3 M QJ c e tn QJ \ -H M-l > )H ax: u ■H -H - x: Vl Pi ^ it: o Vl 0) ■p QJ 73 x: C +J 0 O >1 ^ <^ & x: 3 £ -a o U-l tn ^:?v^Qj Pi tn U 0 AJ QJ E P> J-t 4-> QJ «c QJ M CO 0 £0 0 CP 0 c QJ U '^-c .^^ >iP 0 a: to Vl C 0) QJ E Pi 0)^ '^ -,::-- to CJ u H QJ 01 >i . VI ■H V4 e -H 3 ^ rH w s: ^ PI TJ iH a D^ 0 Pi 73 Q) +J X) Q >.XJ 0 O OJ in -rl ' . -H C 1 ^^Tfi-Q 10 CO U H tu O rH u QJ CO i tn 0 to 0 u 0 > to to 4J 0 QJ • P QJ tl) ,-i -H C Cr* a C Pi Pi ti c C arH 0 c 0 -H Q Ti 0 -H ■H +J -H TJ ■H T) CU QJ Pt c Ja CT>'Cl C P p) to x: en H -rH c c to (0 Pi to 3 XI c to •H (0 e QJ x: 0 (fl (0 -a -p D^ Pi x: ■P OJ c ■H a» QJ 0 T3 W C 0 P> Xi -H P c u a QJ Vl -H Vl -H QJ to -H Pi CQ QJ -P C QJ arH 0 ■H > 3 to TJ tn •H 0 -H to X) -H 0 H • C 0) -0 ^ -H r-, 0 to x: Pi 3 > ^ QJ 3 to CT QJ 4-) Q Vl to c Pi QJ U M Pi fO C 'H 3 2-0 C ■H tn tn QJ j:: 0 <0 0 O C P 0) to < -p -H 4J D (0 0) U ■P Q) ■P r-l P XI tn M P Pi 0 M 0) rH -H O •H CO 3 to e T) J3 -H TJ tu M P 4-3 O T) 1-1 TI QJ p x: CJ P s Pi (0 fO X < ^ M JJ Q 14-1 O 2 VI u D C . H 0 e tp o to c OP"-* pi in -a O C QJ 'O X XI 10 C OJ o cnta lO ■ OJ S H M c/] J 3. : 0) 0) • 1-H x: O 0) o o c x: c c -t-J O X> fn += c: O ceo oco) cm ■H:3x:rHaa)> -OrH OHMB>a)>,oci3 >>aiHHHajt-iti E H ro 0) ? t-l 0^+^ >iH ^^ > MP- m rH c ajtO'HCw C3j-tSo o C w ei3 >i '-^ +^ Ha)C>C-H>'H«)CUQj ■PB0MrtfH3CC=l ■HdjmQ; 0U30W O +J 0) C o '^ Vs bOH a PhOC^ o a w x; XI a -t-' " « i; 4-* to -H o m U T3 Vi i-i o o o o o (u x; <*H art +J +J. -H OJ O r-i > n X) x: 4-> 3 o c • ■ a > S 0) >i-c3 - _ _ WOCt-iCWi-HP. '^^ oj HOjaJwo. ojo c tiD x: tu X 3 +^GCi:n> f ' 0) oj 0) a)x:o -oox:> w rj > iJ o CO &-*-> M art 0) bo 0) rtQ)x:c(UHCo -co OX3 o^ia-^+Jcoc 0) O □Q'O rt"0 a rtXl (DrHO rH *J c X bca 3Hajx:BrtBx: oPhCXo oara-p x:aJ0J3wMrt><'H no'oa*j:'*-<'aaj3 0) +J +ja>* cu H s-PxJ X» 'H xlf-tcCCr-H rt 0 sco3cd a 0 M H ort(i)H3 a-a P ocutn rH n i-( i-H 3 -p ftrH 0 HGcort x!x:=a rt rH H BGO.tfl5O-Prt(U4->C0OO bO 0) -3rtx: +^C0 rHCC-PCOCUM hO-P 3 C03-P ajrtaJ-3rt o-CCto t^ rt i x: dJ w 13 ■t-> QJ H O+^-H a 0) -PX: BGQ)CO H^ OJ to B M x:rt-ciGx:rH+^rtSa)3o^i > rtoj co-Pi-p op-^a> -0 +j 0 f-lC+^ C04->rt Irtr-lC -H-C Q) CO -p m '■M+JOC^QJ(l)OQ)S 1 COrH f-l^4-> m Qj rt oj c:oco-H Gnt-iajf-iaiaiS-iajaj+J'H 0 ^1 x: 0. :3 +J T3 ■pOTB-HtH-a ■rHx:tpM Q-^ 0 c to 0 0 m c njQJ COrt.-HQJ'OS oB ioa>MQj ca ^( rt -a iO 0, 0) c HrtoG. o-p(y"ouoto f-i>»a a oj en ■PBC 1-13 rHCUO 3»(l)^S^ Q) ^ m-H 'nH 4->+JM-P>,+Jrt-— ^XjrHO) x: c 0) 0 (i)-p(Dcn-Hcort+^D^ ti03o(uart-p co> •iHCx:-'Ha>>Moa;cji>»3 ■p(uti3ri^o-p^3a)(ux:^coc-PG G x: H xx:cp Mrtcj-H -(J 0 x: 0 ■H +J +J -p -p tn U ai a CDC ojnJ+^+JCOiV'Qj.H+JCi, rt 0 > p 1) mocnx; rtoo 3C (ot3 dj (u-POrHco c-Ptio d+^nJotu a f-i 0 -a P. cu +J M QJ f^ximi +J3tH-ortrtCaJ-Pco-p+^G D- 0) x! ^ a (D^-'-G-H 0 ^1 ^1 0 H 3 CO oj 3 x: n c 01 0 CO COSO-OOOJCO 3Gf-iO-H3C0 0 3 0) rH fUjD3HrHbOtUC C CO™ oBx!-Hrtcoort a> qj-p to V^ C 0) CO +J 1) 0 x: CO CO Jh W -P QJ Q> H-ciiD(U03i3 rt3B^BG>'-iO OJBJ^O C^^^'l'B rHO-HtH p-a p -p 3 -p rt >5 1^ >. H 0 rt G 0) OJ w x; -p +^ CO a) x:>-'H o>oxi>-H -13 ^ . . ft) OJ ■H rHx:o(u(-.bo5 tio-p+j+^x:cotHCa>aj rt 4^6034^0101)1 +» >i J=i a> ecu OOCDrH^lia) QJ Q) Q) 10 +^ cfl x: HX)xJo(UG2:co t^—xi+J xlx; t ---S-P Ci r-i CO *H (D+J^ioj o-'-t>inoa-p^» .P+J 1 3 H H C03 0 JDHaE +^ rt ^ GO B rHM3T3rt+^^.mp ft-p = w ' 0) G tuD rt v 3rtto3 rH 0 0) tn r4a'a>(Ha>-H3+^ >>'0 CO cj CO a Tl T^ : Q. H c a> 0 rHpHtOrtCOOXOCCOM -H HnH ■Hrt ort(oa)03'ucoc+^'u+^ ' 0 -P -c Vl t 0) 0 QJ cc oox:rtrtWHO : 0, aj JD a X3 ^3i-i3j=i^t^-'3ii)(D3+^xioa)>iC Cl u *J i4 o c £ ^ D 3 O = 3 3 = & ■ rt u C C 4-1 W C JJ 1-1 a< ^ — . tj-. u OO 31 u tn u Xi g 4-) ^ 0 « ■0 ■H M M Oi ■0 •0 (U m v^ 0 a (U o 3 3 (d c x: u 0 Q) Lj . x; a m •a M j: 1) 13 1-1 u (1> 0) a> 41 > C 3 01 > § j= a u (3 rt XI jr H 0 ffl JZ u M d 1-1 , n M 0 d OP 3 4-t u u -H d d 0) x: « 3 0) 3 nJ ■0 X) JZ w t-i 3 c M d u d 3 ■H ■H n) [b 0 OJ lU 0 w •H 0 a. X u OJ U-l 0 -H 0 0 i-t -H (U c rt ■H c 3 TD 1> crt St »-> c c 0) 1-1 d 0 00 0 *-• <-l to O Vj t3 1-1 ■ °°|- •a d Cm x: (J Q) u O t-i oj u D a. tJa^4Jn]3tnwc C -H O U d Qi 0) t-i T9 D- C M 0) (0 0) -a -H o « 1-1 •-< -H di 0) nl XI wOi-iX-H^njt-i O CLh iJ (j O- ^ 0) B -a MB > domxiouod o nJ a (U t3 iH dj a. -H Q. 4) -u w w C >^ O 01 ^ •H •0 -a 3 x: CT] j: d (U 3 a> ra 4-1 lU i-i n) J= 6 rt (U a 0 x: Cl> m ■H D. > U 1 3 l-i ^ d 0 a; rt ra 0 XI >s Q. u -H j:: 0) o x: x: d d a 60 m o o -i-i *j d o QJ JZ u ^ -H u) j: -a * .§ " d (U 0) -r^ 4J x: 1-1 41 0 C 4-t >. 41 M t3 « 4J to > )-• 3 > x: ^ 3 0) 1 0) 00 41 n] M t-l 0 0 tfi kJ 0 --i 01 u T^ 0) )-i d I- ™ x: x: tfl u 60 0. c )-i 0) u -H -< U) 0 •a d 4-) XI 0) 3 n 4) (fl 4> rt X 41 « 4J 0 x: ■H >> H d x: n] M U u u fH m iH 0 0 x: T^ 0) 0. m Cw u t-l d m M D. -H V ■H 0 •^ 0 3 x: JZ >. 4J m 3 a d M 0) G d 1 3 0 .9 4; 0 a V 0 g ■a 3 0 0) 60 41 x: 4J «J 4) d 4) Vi )-• w 0 rt 0 0 U a. 41 t-l x: aj > rt « u 41 u 4.* 4-) d 41 41 (d VI a 0 x: 4) 1 41 0 QJ -H > > a. tn 3 4-1 4) 0] s d (d ^ 00 x: u i-i i^ flj > (U ■a 0 ■H 0 0 0 4> ^ >> ij -O 4) O W 4) OJ d TS Ij O , d u) o u 4J 0) > d -a o x: -H 3 c 41 41 ^j d 3 3 0 ■H 4J U a QJ d ffl x: 4) M 4.) XI 4) 41 4-1 41 4J CO 3 (0 0 a jj 3 d s V 4> nj x: d • S o 4J o -a U3 -H 4J C U) 41 60 QJ 41 i-> ^ d 4) 41 U-4 j2 4> O O U-i 41 4> 4J M tJ > > 00 4) ; • 0) . I 4-1 H jr c fd V ^ ';5 t-> U-l ^ 4J 0 cl 3 CM ^ QJ a 41 4J > o V' 4-1 0 3 T3 0) 0 4) >.x: 0 x> 4J XI >^ to X) 4) a to > to 0 T) 4) 0 XJ •0 a, a a CO 41 u -H ^ cd (d o; 41 c W axi x: 4) « u 4> 4-1 4) 0 V4 XI a u (0 4J 3 d (d 1-1 (d 0 4) x: c « x: XI > 4) 4) 4-1 (S 41 B 0 -r< 41 )-• Q> 0 3 l-l -H 0 d CO 0 4> d 0 u w x: 01 •H trt 4) a> l-l a: 0) 1 > (d 1-1 u ra ■H 0 01 rt x: CO c u to a to O C l-l : ^ Q D l-l o o •-» I ^^ : > C 00 o d Ifl 0 4J ■ 4) 0 41 (U s d l-l 1-1 t/1 u ■H oi 0 x: d 0 ^ > I" Q W d Q 0 u d 0 c x: 3 u E UJ a 0 4J >» § 4J d •~i C l-l s IM d 0 0 > 0 0 QJ U 4) B 00 u Ih DO 41 3 d E S d QJ 4) 3 l-l -H •0 s •H to x: x: ■H 0. (0 to to 3 60 l-l 1-1 41 (d 41 (d 0 41 0 Q 3 Q 3 3 Od 0 a. to 41 t3 D. Cd S 41 (d to o 41 -H O t3 I > 4J 3 1-1 C O 01 4-1 3 CJ 14-1 C tJ 4-1 O M tfi i QJ d 41 4-1 CO o to X 41 1-" M -a J* H E "3 o d 1- t* x: x: Q) IT) a cd °-fi I** 1-1 QJ CO 00 (d CO CO • U «4-l U) •0 Cd l-l >-i d a. 3 41 (U to 4J 4J 4J QJ X (0 (d a 41 l-l -H >sTI x: 3 41 41 5 d i 3 QJ d *-» 4) o o c a. o ^ V^ «-> 1-1 1-1 4) 00 (fl 41 XP Cd - d Cd ca 3 a. u >. 4J d 0 0 Cd 0) 1 41 0 u >. x: CO 41 3 x: TD X) •-1 4-J QJ > 4-1 U-l 3 d TJ l-l 0 0 u QJ 41 d 4) 0 Ul E l-l CO X Cd a JZ cd QJ cd 0 3 OJ 4-1 41 u 0] 0 CO 0) 3 1-1 u QJ 3 •-* Cd 3 4-1 ^ 0 l-l 0 4-1 U-l u 41 P-.X1 cn n QJ XJ 0 d x: cd d XI QJ CO 4J 4J ■t-l Ql 0 IJ 4) x: to I-. J2 1-1 to 4-1 -3 3 CO c ■H 4-1 s x: 41 u- 4) l-l u 3 >. to .ii 41 e cd cd CO X3 tj > Li 0. 41 4) td a 0 •a x: a. 41 3 0 G to d 41 0 u 1-1 d c QJ 0 0 <4-l N 0 tM 41 d to 0 ^4 u 0 tJ 0 cd 3 ^ 41 0 d a 1 QJ 3 a c s . e X) 3 u 0 cd 0 G d § ■-I 0 Cd Hi T3 to ■3 01 4-1 X a x: QJ ^ d 0 CTv i-> 3 60 >. to 0 41 0 ■H cd d to 0 1^ 0 u 3 ■H X) •-1 41 u QJ 0 tn 41 XI XI >. (0 IH x: x: d u ■H 0 a l-l (d d 4) 41 Vd ■T3 1-1 td 4) td x: 0 0) 41 t- d § > a 60 3 >. oc 3 u d 0 td 0 cd ■0 41 Ql u 41 (d c x: c 1-1 cd cfl Cd u 0 cd T-t 41 a > £ to x: 01 01 4-1 CO G to 4.1 cd 4J 00 3 41 Cd 41 to d 60 0 1-1 U-l to l-l (d 41 SI 0 td 0 CO x: >. 0 0 -3 tj x: 1 d d (d 00 3 QJ u 0 >> 0 01 G CA !■< >. a 1-1 4-1 u ■H QJ 41 J£ 41 x: X) 4-1 X) XJ JZ u a i 4-1 C c td u M t-l 0 to X CO QJ XI Cd n G TJ c 0 CO a. c a Cd to 41 u QJ X Ta QJ 41 XI a > >. 60 3 J= .H ^ 3 0 3 0 G l-l CO 0 C 01 3 3 S 3 T3 to Q> d u 0 0) QJ ui! QJ -a 41 d -a 0) 41 u a u » tP 4J -H rt 0 J= Q. • M m -P c -u (0 n rj a< ■H ra o Q) ffl 01 j:: (J^ E 01 -^ 4J lA tP C H JZ m IJ VI -H 4J JJ to Jd C o j:: 0) V to 0) m tJ w o ■a 3 ---I 4J 0) fO o • c o H a -c o -u to lO 0) V4 "O 4-> to U M 01 01 -^ M ■■HCnlO>eO(CT«l C.H q a; w 3 o 4-t *j J^ a. Vt fl 0) •o 0 m C o E u E tP rH Vi 0 u 0) 0) ■0 3 3 p c •H 01 T3 01 0, 4-1 (V c 0 3 u 4J (U •H '0 Q> Oi 3 to c O. p to OJ >i OJ 01 Oi E Td VI Q, J= 3 3 r-t > iw to (0 0 01 0) tn H ni 0 01 ■H -H ^ X o> 3 Vi > 3 >i c -H 3 4-t -0 3 01 a 1) 01 01 >. C 10 M XI to 01 o o> Q. c le E u u Ul Xl C m a, Ti c to 01 0 c 0 01 U) p to o C •fH VI a 01 lb ^ IN CP 3 ja in m to tn 0 m n VD Cn ja o to (0 p 0 q E 0) o^ 0) Vi 01 H 10 Vi 0 a 41 x: HI a* 4-) 3 3 Qt to c u-< U) o c 4-1 to 4-1 01 > 0 o ■d 4-1 01 01 4-t ■H U c j:: ■H r-t Cu v< o to E 4-1 -H E 01 Q> V VI p 0 nj 0 ■H 3 > 01 (B ■H 13 to o E tn a ■a —1 Vj C p D> v< C S o c c 4-1 w 0 ITJ j<: to o Id 01 c to 01 q o j:; 3 JJ o o c ^ •H -a 01 *J ff (fl 4-1 01 tn (N T) m 4-) VI ja c -H to Q) a> ■0 c T OJ m x: mH (T. 0) n c c o (U 4J tn Vi O ■a .^ x: (0 (0 ■H 01 o 0) tn tn m x: ■H TJ o 0) P 1 Vi > E 3 3 CJ 0) 3 c 1 Ot 01 0 to •0 O 4J <0 q 01 >, 13 1 U-l -H n 01 3 •*J to s to P c U-l -C X 0 fl 3 ■H c E •H 0 Tl ■H 4J tp 13 0 C 4-1 tP >. -a o C TJ o; •H 3 0 q 3 (0 Vi 0) c to > <0 01 > 4-) 0) B tP to x: 3 tn 01 3 V to a> p •H 0 CT> CQ o tp -r4 C n) <0 01 q u U m 01 c q x: c E XI o to 10 Qi 3 -H 4J to >1 o 01 1 CP <44 0) q (0 q x: IN ■H CO n p — to o E 0 0) O 4J tP --4 E Qi O ^ I P O to 'H 4J 0 o c o ' 4J O O U -H >i -H tn -H to P o a: to ex O V n) £ El rt >i *J JZ -H > -H a 01 01 J= E i: tP fH (0 P c (0 0 \ 0) .H . 0^ tn 01 - nj TD q > o to 01 H 01 > -O -rt -CM T) to >, 3 nj O to J3 . 01 N O 0) \ -H (0 q -^ "4-1 c m ' 4JpqEotp'Oo-Hto to q 0) 0- x: 4J ni -H u tp o ^ to > q Vj -d p c 3 T) tfl o o q ipoiooatp-HnHto qx:-px:eco-Hto .<3ifl(n--H-H«3a o < « = 5 : tP -. 4-1 13 4J ■H in a< P 10 01 P M J3 4J M-l U] (0 01 TD 3 >i C 14 c ■H to CJ q 01 XI J= 01 E 0 n VI ^ 3 j:: O Xl E U-l to to to 01 Q 0) cn 4-" 01 VI IM (0 01 x: « 3 >i CO 0 4J ■H TJ Oi Vi Pi nj 01 o 0) (0 0 J= 3 iw c ■a 0) O JS 4J c VI .-4 M a 0) 01 >M 01 o 4.1 o m 0 to x: tp 01 E 4-1 U-l HI c 01 q > J3 p 0) q u 01 to u 4J u q q M ■H H —I -p-t c P x: •H q Vi a ■H 1 H 01 01 <0 VI P 0) nj a u > VI c rH x: XI "O 4-1 0 I tn >A ti> VI I tn to x: 01 . ID x: p x: u o q 44 01 01 u a -H o Vl 01 01 ■H x: ^ rO E x: c H ■H E 4J V4 Vl E in 0) O O ■V OJ u Vi Vi q IP - to 0 •H .-( n o o >i o E C q j= -a 0 01 3 3 01 ' tP w q tn to q 3 0) *J j^ .r4 P nH q CP (0 « 01 c C E j:: E O ' < tn > 10 ■H M *J w U ITJ E 4-1 a c d M 0 U >i 01 3 x: VI to U a E q u a 0 E i-i o ■H E u 0 x: c M o u 01 3 Vl u to 4J c 3 to -H Vl to 0 tn 4-1 (0 x: 0 T3 'H c 01 c 4-> ip 4J 0 Vl c < 4J o -H 01 Vl q to 01 P p E 0 0 x: to to c c 0 -H 4-1 > 01 0 P OJ Vl u u c 3 (0 x; 01 01 .H 0 O 4-1 4-1 CO Vl > -H >• ■H ■'-1 XI c 4J > Vl cn 0 >1 w (0 .i« c a c E > c -H < 01 p Vl <0 T) E 4-' q 01 j:: to 0 q o to Eh 3 01 10 01 a M c O p Vl E r> Q 0 >. to •a 0) o to Vl 4J 01 CD a 0 to x: 01 r- 01 u P Eh x: (P 10 > 01 w —I to QJ 10 01 OJ > -t TJ 01 O H O J > ^ 4-1 ID OJ o q c E fl 0) m q 10 Vl -H 3 iJ O q u CO o Stf O O to 10 I "O to M tT" E O I 01 >i to -I QJ XI -H nJ . t OJ TJ VI x: IP c 4J Xl30ia'+4P3'D t 4J E « H a -v "d ■ x: 01 o q o U O QJ 01 01 4-1 4J P (0 1J TI CO tD ■H -H a 01 01 3 0 4J E 0) 0) Vl 01 >t E 10 M J o 4J 01 D> 0) a d J= rs] q x: >u 01 01 *J — ■H Eh 0 Vl 4J 01 tn q E 0 0) r-t Vl o — I ja q ID q 01 Vl p •H tP J= o c >. q q Vl oi'H q OJ o n m at ta jc 111 01 3 4J to > m Vl p cn Vl o 01 a w to p o( >, S 01 -0 E E -d O tu E uj to to Vl H q CO H 0) tP 4-1 01 4-> . x: o o -^ 01 o > tH 01 u ID x: u o (J s: Z '■. > < • 1 1 1 * U Is < i" 05? 01 -H OO (U . OJ • C u 3 J3 0) 4J 3 ■H C O yj 4-1 a OJ n D j: OJ > 0) ^•o" 4-t CU 4J -H X JJ W nj 4J 41 W OJ 3 m C OO 4J ca -H 4-1 c a o O X X n) ■H (U to (U (U ij B £ > o l~l .o ■H O J= o 4J U) hJ n) *-t B - rt D. to m ^ (0 !U U 0) >-l w oj n -C -H ■H Ji c > Q) 4J 0) o I]> u a> c. u ^ j: 3 U cu B h f-i 4j O 3 (0 ^ « X o 4J S l-l >-> -§ Cfi w 01 i~> h-i o c u 4J T3 Q> OJ O B o> 0 a 01 If) T^ 0) J= § -^ > O 4J 0) 4J c e VI Q. U ^ o o m 4-1 O 1- on 4J •H U XI D. U CO (0 3 *j ^ ^5 M c 3 U QJ 4J O -i-l 01 CO -H CO 4-1 OJ a> j= J= e " ., J3 J3 -U >. u to o a; OJ 0) U 4-1 B --1 t-« rt o i-i a; 0) 4-1 cj -H 4J £ to 4-1 C iJ 4-t to ft I-l o U 03 C *J U O 1-) X rH iH 4J -a 00 JS V -rt -ri to ot rH -a B o V n u v J= -H O ^ CO 4-1 U U 4-t 4-1 U CO h 4J CO tu o h tM O > (u a> c 4-1 3 T-i 3 (0 B C 4J 8§ a. e 3 -H (0 ll> 4J )-• J3 to s^: 41 C 0) W "3 ,c o ki a -H 4-1 u (il to "i-. X c C 4-1 (0 M o OJ tt-i U 4) I Ti CO I B ^ U u a> OJ 3 o c u 4J 1.1 41 CO 01 > OJ :=i • 4J <4-. C 4) to t U (U O M to to c o « a u CO 4) to 13 W to .n 4) 4-1 k4 Li .JT Vi tn 4J Li 01 4-1 4) to 4J 4-1 -H a. I-' CO c t. o. w x < o 5 CO o T3 - H §ji 4) V4 X CO O 4J Li '3 3 4J .-H .-I 01 ^ Q> .-H H I -rt o c to O L. >, f( . > X CO 4J a> CO Li 4-1 s s I D.O I Q tNj : U-l ; X 0) C O 4) C 4J B 4J » J3 .a 3 c CO li-l f* 4-1 n 4J c J3 01 ■H c to to 3 4J to o 00 4-1 c CO o p^ to CO 01 C OJ J= CO 4-1 c 3 >. QJ c r-t Z ai a rH C to ■H u tr tj CO c Ji o CD c B Li £ 01 CO e o to c CO "D 41 0) 3 Li u x: 4J ■H -3 o a> Li 4J 0 O. 4J B 3 4-1 U-l 0 «J V to to X -H o O. 4.> 41 CO to < TJ to en c O CO to CO T3 3 4J u Li O -H u-< 4J OJ c o X to Li .H x: C CO O Li C Ot to ai fO 3 i-" 3 3 to ■H -a <: 3 o •D I-l 3 to D.= a: ■Si u r- n) o^ u a>tflcDO itOC'MCtnOJO'-tiO-^r- 'OOO3Hj::C4-i0t4\ • fO U "0 -H 4-1 u c ^ to c rS (0 3 OJ OJ OJ c o *-» ■ n] lO = 0 tn OJ 3 a o 3 c « 4J ■H 4) a 0 Li 4J 0 a> o tP P 0 4) tn TI CO c a; C J= r-t VO c -H M ■H OJ 3 4) -H P 0) o o lU 4-) tp m 0> V4 J= 4-1 TD > -«• fl c 0) O « 4-1 B OJ C W V c X c Id m c h o to 0 4J -0 tr- 4> o U (0 u 10 T3 O <*-l t) O ax o ■P c o 4J 4J OJ O 0) 4-1 iH o 3 o to 0 0) o m 4J o OJ Li u TJ 4J o p ■d ■H 3 E nJ o O 3 Q >1 C o ■3 41 X to 4-1 in c C TJ k4 w IH 0 fN Tl o c p p OJ >. to 0) 0) 4-1 4J 0) G) (0 (0 P o •D c V4 X ■p 4-1 p to 41 OJ 0 o tn CO p to 41 t^ 4J c XI > 1 E o 0) 0 aj QJ > ■H to to OJ 1 O 3 a E P U •-t a> OJ a o u to 0) rr OJ to to « (tJ M (tj p o ■H CP k4 O -H X 01 D^ -H i-i c X c to 41 c o X V4 to (J> p 3 5 —4 ■P o m *J ■a X rtJ U) 0 c 0 u 0 4J M 01 (0 OJ 3 3 ■H c M in n3 o to p c c -H t7> p Tl ^ 0) 01 (M 4) a -H 0 3 <0 3 01 41 E 4> > >i OJ ja -H c E o Li -H 3 01 —t E -o M 0) to E p TJ c OJ to to T3 r-i 0 H n X nj u o nJ C o X fl X to 41 C < o > x: Eh 3 — to u Z > P •D to a u u u o a. 4) Li tJ o -H > ^ < CJ Ll X 0) to m C CP 01 X >i 0 Q ■H 4-J >u -H 01 H 1 ■H 01 ■H o u ^ X r- p ■-H >, 10 o^ cr> o o in o ^ C a a. E 4J to a: in OJ 01 c tn X X 4J <4-l m 41 p p ■W to > -H 1-5 > — U p 3 U u n) OJ to 0) OJ to X T) 0 X > TJ -H ■H p P o c tr ^ to c c 0 JJ rH c to OJ OJ 3 O -H o 3 u nj OJ V4 u a a b* — •-I tJ Jd 0 >1 •o tn OJ c o OJ c U T3 QJ c ■H X 3 E <0 ■;^ p U 0 o c E 0 p Ll _' a >1 0 U o +J 3 c to a c « E o 4J -H 41 E 4) C 3 4J c (0 E ^ to 4) OJ OJ 0) P O L< E U X tn P to u o a 01 X p to m X X O X 4J 0) 4J J£ p 10 0 OJ to X -H 4J fH X u-l c a p X C p 3 to c 4-1 ■H o 4J x: X (M 3 e: m c u u 0 c c (0 OJ 0) 4J 0) e <0 OJ X E 3) a >1 -H 0 o u p OJ a 4J 4-1 o Ll to o P 01 P 4J 3 c -H u 0 3 w ■H 3 0) Ll X X — cr X p to u- 01 to u x: 01 4J O -H r-f s: o j-i -t-i in a C -U -H (0 . ■<-! 0 O -tJ c O i^ o J-) c CL--H rH JJ >, 0 0 O -OJ'O'O'-Hl^J-i ^ >,4J fO fD fO H.-'-t C-l>£i-HUUCLUC C'OT)Xi--*aoa) D^ 0] nJ Uj (U c m o TJ x: >i 0) (0 U J-i i-> T) MJ (0 (0 u c o > CU fO 01 -r-i+J C «4-i x: XI in o 0 ■*-> O -H w q; 4J 01 -n x: u - . « „ . , X! C QJ -P 0) X > -H fd fO XI -P xj-Hu-i'ow'otD TJ x:o C O-H CU-H fC E >,.H UIO Cr>l-i Ctfl D>QJ---Hcncao»o -Hn3J-iTjTt> >CUOOU-P -Mxo--io(D)-i xux:c tflP>DT3JjO0J-lJ tf)(U>i a QJODd TJ --^ 'O'-l 3-pjJx:'H mp4Jrji/] -H-H cranji'5uiMa)r-(poc>> ox: xo>ujaooo)n3 COS'-OI ■-* C -H-H CD ■HX OJ WQJ-H- -PUX QJi-iEQJ+JtO 'O0(0-H = 0-HJ3[nEx)C4JC«-(T3 tn *^-' cp QJ (D ^ to Cr-H QJ too oovjx:3m cr>-'-< 4J j-t o ux 1-.-UO-MOCCV) ex (/) -H O -1-* 3 -H -H O O C7> XTJna^jl-i-HtiioEV-i OQ) tr-Hoao orj OXWS ■H^in4J4Jcy4-)aioou XOO3f0X:n3ljXU3'4-iO = incc:u-i3+JSfC+J(i3cy]OXl C-HXM (00-PCC O-PHlJ >l-iOO ■H U 4J « (0 T) 1-1 *^ U COOOlCl- -H-H fOrt •OD-HX: u> O-POJCtr-P-PD^UC O C M -H -H -H C (0 Q) .-I<[J4-) CnH4J-H Q C a -H 03 +J CPT) o c c . fO -H (d o c W tTJ l o --< a c o 1 O O O rO jC x: «-. I 4-) +J JJ 00 o e ) m X (13 : -p -D W V^ -P .H C -P c c 0 C CU -H ■H o > fd -P U 0) E 03 U U d) CU (D 01 x: X a-P u-i -p O O I tn o 3 in : 01 iH 01 fO tnXJ O XI W -H fO -H tl I ^ a-P o T3 -H rO >, M o u c a--i 0 ■P n: O £ T) X ■H U-l O O en C T3 U -P 3 >, 0) C XI M C -H 3 • O O O O -P X -P -H O TJ C -p u in XI c o u N u ns o -H o a E 1-1 -P O E o c o O X -H -n 0 -P D O +J 0 tl X X 4-) cri 01 -P ^ >, o ra c -p -H .-H O fO X QJ 01 S o o >i 3 O -I s +J - O TJ C C O Vj o 0 W fO ! ■ OJ -P -I ■poo>c:o-pxc S-iM^hQJ'C-HC-P<1 a 01 E ui n3 ^ -H (0 1 Cr-P in QJ o c Tl 3 1-1 < o 01 E - O X -P X JJ O W 4J O in tj3 O O -O (0 O 0) 3 OJ OJ ; 3 -P I w O : 0 -a in c O fTJ CT-H C (13 ■H E fl E « -H U -H C J-) Cr> O O QJ OJ U (0 0 -a X 01 . 3 in fT3 0) o C X >i ■P (0 ^ a 0 o X to CO >> M -P ■H C -P 4J -H :3 in u a 0 H 13 O i^ .-H ^1 4J O O -H c ■P > QJ 0) C O D- E ^ XJ tn 3 u 0 0 MO -P h >i CO O (0 t^+J O XI M ■ O fO -H E > o (0 , I *J 0 Ofd'UOUOOO in 0 y-i -H ra *j c n u -H O -H 1 u X O -P o -P 0 a 0 O TJ -P M in XI -H >i4J u cn fO E X -P c u D -u u . i: E W 0 -P -P 3 1-1 o o in u 0 tj 0 X CO fU 0 (0 U E U -P C U (D tp O O CM E M -P MO^rOtOtDfOU ■H w -H o 0 n] C 3 X ^ -P M C X 0 -p M cn < 0U(i3ClTiC(-l-H X C (0 ? -H 4J ifi -Ji < -H 0 0 w 0 -p o ro x: •-( 0) 0 U-l E -H 4-» X > ^ ^J 4J -H X M -P 0 O cn -P ■p ^ C 0 3 -P c X X in fl3 — QJ 0 -H . X XI QJ O 'D -P -P -H .-( • in CrO X Z U 0 C £ QJ -HE W 0 113 3 . OJ * 4J ■HOinoSEOM'C U fi3 DOC M cn 0 -p 0 j:: o 3 0 > U-i S-l 0 -0 n3 m (X CpO -P XI r-H > o 0 TJ a-H M E c E m c — e u 0 C Vj Qi -P (0 fO 0 O -H tj* o X U 01 X -H fO ftj tJi CPE U-H.-I C U-E o O -P X NO C 4-> fO -H C cn 0 H TJ -H 1-1 ■H 0 -P U O --I -H >, 4J 4J X f-t C >>3fO'DfC'-H4J ■H C 1 OJ 3 fO iH ■H -H X CU X rj fO QJ 1 X 0 CP (0 .-H in cn 2 0 o (13 E 10 in 4J 3 )-i -H C - cn 0 CO J-» u O O O 0 o 0 0 W C -H .-t E-. -H M M u ^ o o X X M 'H -O rtJ U •p •m a u 0 3 >i a -POW-POC-HO ■H 3 M in U Q U XI 0 J-) -H u 0 O 0 0 - o i-( a (0 -H u -p OTJxwEinxtP OEOMO-PMU a 3 -p 0 0 TD cn c C 0 l-i C -P >, 4J o tr- 3 3 -H OfdCUOOrtIC 4-) d -o C 0 '0 tr •H -H X -P < (0 ^ C -H MO 4-> C O -P C C o 3 > O O N +J (0 0 Tl X M 0 QJ 1^ in -H .-) j:: -H 3 XI -H O -P iw X H 0 -D J XI -P -H J3 C X TJ 0 O C -P 0 O (0 0 fO o cn c - X 0 u M C C O -H * EfCOcncn+J-HX -H tr M o o c 0 in E ^ E C M -P JJ 4-> 0 o in M o c 0 E 0 o c-^ u -a > TJ o a 0 U O 0 -H XI •■ O 3 l^ 0 C vj cn -H 0 -P U -P E "3 > (0 o cn (13 0 • -p u o (d o o c -p Jj •X) IM 0 (0 OJ in 1-1 c E i-i 0 c c 01 cn u c M -H (0 U 0 < y-i X (13 >i C CM 3 0 tr 0 T) 0 O C 0 O CP Oy-(ccxoo-H o -a to fO M c ^0-H4JXXUXX^ OW-H O X 4-> 4-> C ■p 4J 3 X X iM -H X cn 01 cn 73 -H 3 0 CT>-p c in in ■PX-H C C«-i 0^ 0 X -H -H 0 TJ -H ■ H 4J ^M nj -H 0 O 0 in in X 3 U J3 "4-1 X in •H O in p^ in -p (13 4J ■p u -P (0 c fO C 4-) O CL< 10 E E OJ -u E E ro fl c > u 0 n m ^1 0) CU -H in lU > c Q C 0 u o 4J U lU O 0 M O c o cn ra M O O u 0 3 -P fO iP o E 0 o u o in U-4 4J U-l X 'H M M 2 X ' o 0 C -H ■p CT a o .-t 4J O M < -n o 3 MO in 1 Tl U 0 O U-l 4J 0 -P o U-( JJ C 4-t N u o ra cp c 3 0 fO rO O ■H 0 M X O-H cn X -P cn TJ 4J ra -p M cn ■P 3 - (0 -P u 4J u o c ■H C O «P u u ra ra T) X ra O E c-H -H 0 a C 0 H U u C E 0 O .-H iw 0 0 C -H •H 0 O cn cn o H-i O o > M M U-( in tj' 4-> T) O -r-i X ■H 0 • 0) ■H a ra - 4-1 a> 0 in E C -P < U ■H C 0 >. - O O e >. (13 3 o ^ 3 -P U-l tn 0 O a 0 E -P E cr O 0 ra u 0 M in • -H 0 c ■H 'D X a o W .-( O O X -H > O o o in 0^ o 0 X > u in 3 u O M O M X ^ D. (0 4J fO in (0 1-1 O 3 4-) ra M ra 4-J U H X o 3 ^ 4-) ra -H ■H T3 c 1-1 TJ M 0 -1 3 -P 0 c cr> • M C 0 o O 0 ra U-l to 3 -H o c in O (13 01 i4-( 4J in c c ra X -H o TJ M 0 -H M O M 1+4 4-1 -P 4-1 in u O 3 o in a X 0 0 3 M in M M (13 > E 3 o o X 0 c 0 3 t- J U X • -p 0 ■P U 0 M 0 U-t D^ 0 a-P ra 0 M C " T! in ^ cn 0 O -P u c 3 0 a c in T3 O c u O 0 4J E -H -H 4-1 ra M E (0 >. m X -P -P TJ ra 4-1 X c 3 O ■P to E A-> u ^ c M cn tP o 0 0 U O H o 4-1 U 3 ra c c E M C IM O cn a o U-4 M ra 0 o a 0 -H g 0 -H u 0 in 0 0 ■P X X O -H Tl 0 u-l M 0 u o rH u-l c cn 4-1 Q 4-) ra u o ra S: 0 > o cn 0 *M < ra o J3 E Uh H > o )-t o 0 4-1 QJ ^ ^ >, M c o o 3 -H E 3 X 0 cn t-i ra cr 01 u D ■O M ■P in ra -H 0 ■p M -H ra o c tj* o 3 173 -H ■P CnX E c M -H ■H 4-) 0 0 C X c o ra ■p in -H ra > T! C -p cn ■p o u ra 'H 4-1 E 3 O o QJ c c s: x: -r^ --^ u ^ a O +J ffi X «P -H M 0 ■P 4-1 XI iH H u ■P 0 ■P 0 H o 4-> O 0 o C E cn 4-) 0 ra M U X ■1 IP c a 0 M 3 X -p a 1-1 4-1 :-i m -H " -H U O M-i 3 o ra Q O M in X c 0 C Xi o 4J M (0 c in D> 0 T3 0 E S£ O E 0 c c o U ■H >, 0 X ■H 0 H X 0 4-1 ra in E- CP W H C 4J o C (0 £ o o c C -P 1-t 4-1 M M X E -H 0 ra O X (d 0 o m o 4-1 -H > E r-f > -p o > Tt M u-l CT ■H ■H O 0 H u ra 'H Id u~i C O h^ -0 M tr3 U X in Q 0 H M I) -H > o in n M c o >i o -H o -J XI in 4-1 u -I o c ra •- Tl M 0 o a u o n X TJ U CTTJ O-H J -H C X 3 ■4 o ra 4-1 3 3 5 +J 4J M C n 3 c o o U O O U-. -H ■* £ -P >. c o u r-H O £ O cn 4J -H O C u-l ■ a o -H - D T3 C 0 - 3 C E ■ O.cr »H 01 ' in o X 3 tr O M 0 I < tn o ' O QJ I X tfi O i XI > o 4-1 O 0 in o - a o X M O 3 0 4J 3 I £ 1 O 3 T) - OCX X o u M O .-H 0X3 > 4J O O X D -H ra -H tp cn -H > o ' I > * >-> c 1 o o 4-) U M n in M 4-1 X : o ra 4J I M 4J CP-H * 0 C C 3 I 4J < -H ) C 4-1 TJ » -H o in o O C 4-) O -H C M -H ra M I > TJ O M - >, C M C 4J O 4J o o ra o Tl *■ ra a cn — T) M o O O O O 3 X o O U O 4-) £ I 4-1 a M in u 4J QJ O O O M O X M O 4-1 O X in E 4J O O CP in o cn CT> 3 M TJ in O 3 -(— iTJ -O'-l O M 3 o a o (N 3 0 o o - T) -H > I C 4J O J ra ra ■ : 4-1 M M ^ c X-H.-H O-H CPCn O-H 4-14-1 U (Jit X 4J 4-1 X a-H a o 4J 4J Ora 034->XMC X cp o ra o o I C 4-) C O U M cr. £ ■■ 1 3 O u-l o . 0 4J 0 -H ICO 4-> M tn ra 1 >i a o N I c M -H ra c 3 M o o c ■H U ■H a M 4-> X 3 m O 0 •H tn X TJ O o 3.-H-HC cp04-'-pinora>i : O O C O 4J ■^ 0 XI -H : N M 4-1 M E X O M CJ>-P O-H U-. e o -H -H C X 4J 3 U U ■H M ra X ra 3 4J U-t a-r o o c 'V ^ ■- ou oocpc-Hra4J >MC3UCOWr 030inM-HCC- 3 T) > 4-1 O -H M I E H 4-1 -H O -H 4J U -H X C in rH C CO 4J o in o o o -H E -H M O U XI 3 E X X O 4J tP 4-1 O T3 >i'0 : M >, E ) O o o o 1 JJ 4J Ji T3 in ra 0 -H c 4J O rH X -H -H O . M o in ra E c J -H 4-1 4-> 3 > ra : -H X c ) .H 4J O 0 c < 0 in -H 4J £ 4J C M ra o O 4-> U +j -H ra E-r-, C-H T) 3 ra -o O O 3 x: M -H 1-1 -H U TJ (U -H 0 H 0 u <1> C x: 3 x: >, w M-i a 10 4J c to 4J 0 0 C 0) to 0 0 a •H a 01 JZ p M 4-) P 13 TJ P O (5 -H 4.) tTJ to T] 0) QJ C ■_nj: 3 (0 C c ■H QJ p in (0 p JZ 'H 0 U l-i P w 0 T3 >. P (4-1 10 0 t- P QJ 3 c nJ rH 4J a 01 0 P <0 1 D — . 01 0 E -H m QJ aj in 1 0 ^ a^ D t^ * < • M >: 01 U. 3 •HOP L4 CO c QJ to 0 < J3 4:: r- x: c (0 C TJ 2 >i fO D^ cn c\ u 0 W fl u -u • ■H 4-) C 0 p ca T) 3 a xi M Sj >i P 3 -H 2 ^ fl C QJ u c fd -P 0 i3 >-i u QJ U-i to -H 0 fO i^. a 1^ CTi n] P ■H to 0 > 0 nJ Q) - D. >, OJ }-l QJ p p to >i a c w 0 u CP 0) •-\ q; >, (1) QJ 0 -H .-1 4-3 C M -H -H -o ■H 4-t p x: ip c QJ (0 C > i (y utx: nJ en -H x: •H QJ J= C >i M cn-P 0 .-1 C p C CJ "P -P fO c J3 P 0 -H 0 ■P 0 la 0 ui XI 1-1 (0 iw en 01 3 j:: Lj QJ 0) 0 p x: >i 0 U iw a q; ^ 1 tn x: ■P 4-» P ■-H -H 0 > D ■H P P > P ^ j:: QJ 0 ^ 0 - W 0 3 tn J= XI 0) OJ E E P •4-1 0 P 14-1 a. p (0 tn 1-1 P 0 u 0 C QJ U -H 3 :3 u u QJ 0 > -D 0 x: 0) OJ 4-» M-i <4-l P >-.>P -H .-1 Tj tn N x; 0 :3 i-H ■H P (0 c D c ■H p ja iw 0 QJ Q ■H D 0 D 01 c 2 > cn ui 3 ^ C7iU-( .-( tp fO QJ P P QJ = -*J 0 0 --1 c U QJ P OJ 0 x: ■P E U -H ■H QJ r^ to > x: 4J C 0 -P OJ --H 3 a tn •H -H 4J C (D jj U t4 rd 0 ■H C 0 C D "P 3 S W OJ ^4 I/] 13 0 0 D to 0 0 Qj >i a 0 QJ 4J 0 3 U) u tn u) S > x: > cc: p U) 0 QJ -P (n u c q; cp 0 D < x: x: p 4J 0 -P -H W D TI p 4-) 4-) (0 ■H u-i C 0 c >i >. 1 tn H x: to >^p > 0.73 cn QJ H •— 1 p P 0 4J 01 QJ 3 4J (0 c -H > 0 0 0 0 c 1 £ +J QJ tn ■H ■H tn x: >1 in Q) c c ■H > P j:: (4-1 p c 0 - u 2 —1 •p in 0 >i 0 to 0 P c P £ ■H 3 p 44 £ 0) J= ■P 0) P -H — t 0 2 0 01 0 P x. 0 0 3 ■P 0 OJ > XI u cn 01 X) 01 0 0 0 ■H 10 >ix: 0 ■H 0 0 rH t4H rH -p P E P 4J to 2 in a Qj £ 43 P 0 -H 4:: -H 0 c 2 OJ U-4 rH ■H -f— 1 . rH XI 0 XI H c fd X) u c Id 0 to <0 W P QJ C U 0) ■H tn c p P > P 4J 4-1 p fd 0 44 p «H a4-t to >■ in p 3 14-1 -H 0 44 C 4J 14-. p 0 tn 3 01 -o p x: td •H ■<-( . r-. U 0 to C 01 0 u 0 — 1 4= U 0 0 E td £ ^ C SZ"^ xt 0 0 p tn 0 tn -H in 0 m C c 0 iw 0 p c C ^ QJ XI 0 0 > fd 4: P 0 4:: fd 4-1 0 0 13 QJ 0 0 4= 0 0 • x: td > ■H 44 p b. q; (0 0 *P 0 U-l 0 C 3 0 -p a in ■H 0 in 0

    , U c >i C -H 3 44 p tn ^ 4-1 to fd = c PUP 01 4J •H 0 0 ■P 3 X 0 p H-l -r4 > lU Pl 0 < rH 0 P ■H p tn OJ 3 J LO ^ 01 0 0 f-i 0 QJ ^ P p C P p ■H C P QJ p u td >, 0 •4-1 0 OJ 13 P fd p 0 U -P fd fd 4-t to 0 •X. 2 Uh 4^ 0 fd p X >, • to -H JJ 3 to QJ x: xi 0 QJ to 4-1 to in ■H N — ( c tn u tn to c 3 X) 13 3 >-.M 0 0 4-> 0 p 0 Jj 4J c 0 x: p 0) ■H > c in 01 p in QJ x) •H QJ 4-" ^ in X) fO 3 +J >s — 0) 0 0 0 C -H P 0 cn x: >^ > Wl fO -H 3 QJ P -H P to Qj p 01 x: QJ OJ J3 c u c c fd fd 01 0 U 3 10 QJ p > U 13 POO a to 13 (d P P rH -H to > x: 0 3 to to 3 QJ 3 p a u 0 p (d -H x: c P P 4J ■H 0 43 rH (0 0 p E > C CP fd H ■H 0 0 p p 0 P x: QJ p QJ ■P 0 4-) U" 000 4J to en P 4J 0 3 -H u 14-* x: P ■H (d x: p 0 QJ £ to fd c E < x: 13 3 - tn a^ fd to 0 at P c »P * P E x: c t4-l p P p P 3 ■Has QJ 14-1 p c C 0 to 43 0 4: ■H p U to c 0 13 cn COP = lO ■P 0 -D 0 i-i U M tn 0 tn a X) c 0 0 01 tP to -H -1— > -H QJ 0 c iP 0 OJ p U X) P p 0 0 p to c c fO c c 3-1-1 0 ' P a tn — 1 < u ■ H 0) -H ^ XI XI 3 E ■H 3 M 4-t in 44 >i QJ C 0 C l-H 0 0 -H 1: 0 0 p 0 to 3 to (0 to Cj to QJ x: c ■H C 'H 3 3 P -o 0 0 P 0 0 X 43 P 43 in to 0 XJ c SZ *J 0 tj QJ 0 P -1 c x: H 0 0 0 0 0 Cn a 0 to a Cn to U • rH 4= 0 0 in to ' E 4-) 0 c QJ 4J to to 3 QJ 0 QJ -1 0 p QJ -P P 3 x: c c -H in sz c ■H to £ 0 to 3 p - p 0 13 ■H t-, 0 in 0 cn -H >1 = 1. 0 ^ J3 P C (0 in p u in M p ■H x: 3 .. 4J fd 4:: c P 0 2 -H QJ 0 tn U rH c < E 01 43 <4-> 0 C rH •-1 c j:: to p E to C td -^ -r4 C -P to 0 0 to 0 w u >, in fd 0 ■H H 4-t in c 4: P QJ <0 x: tu P 44 -H rH *o to 4J QJ Qj 3 c a 0 0 QJ . in QJ U p x: XI p -p E X) P E M-i UH C p 0 4-) 10 -n fd 0 cn 0 QJ 4J - u to p p x: C 01 ■H P in 01 0 3 01 • QJ 0 p OJ p 0 c to in 0 C ■H 0) 0 p in ■H X3 p ■H CJ x: P 0 to -H 4J >,-H 0 tJ^Tl QJ P a'o P -H -. a rH X) in LO p XI fd --H td 0 3 XI 0 ■rH P 4:: in 01 X) P 0 >i c 4:: >-> 44 C c to t|H X: rH u ■P C C -P -H QJ 0 0) fd to 3 43 E <4-l P 3 4-1 p fd e > X3 p 0 E 4: c P in a >,rH c 43 01 3 ■H 0 -H Cn P OJ -P -H to C 01 Cn N c 3 QJ td 0 c OJ in c (d OJ 0 OP to ■H 0 p j<; 0 0 X X3 3 0 0 in w XI U P ■H to a ■H JT -H c P QJ -H P 0 ^ P u ■H x: 0 C 0 x: «p £ QJ P C p > E u -H C QJ 0 'H 4J 13 01 QJ (J < xs 0 c P rH in r-l W to XI -H QJ -H ■H p p 0 E p x: a 01 in XI ^-1 0 u -p in 13 30 0 p P [1. 0 3 P 0 3 0 : -H tn i -iJ -1 to P P vA 3 0 >i QJ c c p P 4-> E 0 QJ 0 *J in -H 0 0 tn P tn a 3 < c 0 a-H 01 0 [1. <0 P D p -p C (0 tn x: ^ C -P 0 QJ 0 0 P QJ 01 CP in m ■H tn 4:: 0 13 CU p x: u x: -H QJ 14-* 01 OJ X3 >.x: u IP > 4J D>-H C 4-> p in c P ip 0 CP P fd in P td P Uh 3 c 0 CP a in QJ 2 in 0 in CP in p 44 43 0 c ui tn JD cr OJ E C H to 0 U-i 0 -r-i td 0 p tn r-4 C -H ■p p 0 x: -H E 0 U 13 0 c p 0 0 * ■H a QJ p QJ OJ 0 0 in x: > CbX) to fd c x: in 3 OJ rH OJ — p p ■ HP P 'H 4-t 0 > c c in ■H C OJ fd rP 4-» x: = QJ a E in x: 0 0) u P P U-l p c 0 0 0 E to 3 c 0 c fO XJ a ' c p P fd P [1. 3 0 p X 0 > arH c c ty E ■H p tu xa in c POO 0 P OJ 0) -t E 0 u 0 C 3 XJ 0 OJ td '4H fd 0 p a< 0 u 0 lO fO 1 -H QJ 0 -H -H x: Q, 0 to ■--1 to tn a c 3 a 0 in ■H c > p OJ 0 0 ■H > CJ 0 P tn ■z. P tn E 13 x: 1 3 r— < -H X • XT' in -H P c p p tn 3 QJ OJ 0 10 £ 3 P c P OJ ■H 3 p x: M c tn 13 c to > c 0> ^ P 01 in TJ in tn C QJ ■-H C C a P 0 p ■H X X) ■iH 0 u p XJ -P in rH -H fd 0 p ^- 0 fd QJ 0 fd rH -H 44 ■p QJ j: (0 XI P 01 in QJ QJ U ■HOT] E c 0 0 in c 4-1 P 4J 0 C QJ rH 14H 10 to in x; >i43 3 4:: ■p 43 3 x: fo D > P 1 to E QJ P td -H TJ in c 3 C ^ fO Ui in P P 3 0 P <4-l ■H > xa 0 0 to QJ £ E in P to P in 0 in 13 tr c Q) XI -H p 0) rd to X c 0 3 0 QJ in -r4 in tn H-l p ■H to P 43 13 x: 01 0 C 0 ■H QJ 4:: -H *u 0 *p fo x: E E E - QJ tn 0 P C «4-i X3 01 ■H XI ■nj;: 0 to P QJ to 0 tn C P P rH c 0 rH P E p 0 C XJ 0 X3 in UH rH 0 0 E U 0) -H in x: in T) P 0 0 4-t 0 to p 014*: P p p to u in ■H 3 0 3 uh -h 0 3 E OJ p ^ Q^ 3 D^ E -H QJ to QJ 0 ■H >,U u 0 x: E Oi U in 0 0 ■H OJ 0 u in 4:: 0 01 OJ P QJ X) 0 p OJ fd x: fd en Q) en C -H -H -< ap c p 2 0 3 p - 3 0 0 01 tn 4: U-l >it-- c cn C P 3 P in QJ fd 13 0 x: p 0 > ■p > QJ C ■P ^ CP U Ii- QJ e to 0 to M 13 C to P p x: 44 42 43 QJ 3 c 0 QJ 0 CP N 0 c p QJ in 0 c QJ 0 -H P JJ >i c QJ in T3 -H N p en 0 01 - C -r^ Ji p >; 0 43 0 ■H ■H c in x: x: C -H ■H ■H x: QJ 3 0 tn c P P OJ O-t-H a u E 0 ■H fd in 0 p u in 0 to in x: u cnp 13 0 13 44 0 0 3 p td c 14-1 CP P P x: x: n 0) 3 to Tl > 0) J^ tn -P 01 E 01 c ci. QJ x: a c 0) 0 CP C OJ 0 14-1 3 c tn OJ >, 0 c fd P 4= fd •H c c fO to CP 3 ^u 0 E C > to p QJ fd -P 0 P •H P 0 < 0 x: p 0 p ■H 0 P r-i ^ P ■H P XJ fO >i^P u cnp 0 0 (4H 3 tn 3 0 ^4 E -1 c in C XI en u QJ c in to in -p p (0 44 P cn fd -I C XI ■H OJ 44 rH 0 p tn E 0 10 >i 0 in CO OJ QJ U P P 1 to to 0 (0 P -H c D> -H 4J 4J 0 m 3 CP 3 to in 0 E x: tn c td 0 3 p to c x: P c QJ x: a: -H Q( (0 c 3 -» 4:: ■■-* p 0 P C - in 0) . fd ^p 4J 3 rH 3 H XJ 01 ■H to 0 >i ■r-i to in 0 3 x: 0 ■H 4J p p C C 0 T) P u in QJ p 01 ■H P 10 43 td N 0 i^-j 3 0 ■H in fd cr c xa r-t -H cn P 0 >i N 0 O-H P -H P en u to OJ P -H ■H < to tn T) rH fd 4:: 3 c 10 -1 H r-( -P 0 0 XJ 13 > 0 E 0 rH c to ■H in p 0 -H 0 E P C P ■P P C P c a QJ to > 0 -H -H to CI.P 0 to 0 -t 3 C 0 13 01 0 ax) x: 43 -H c p 0 0 XI c c 0 to 0 Jx: to C 01 -H Id -P 0 P -H ■H 4J u E tn c rH ^ (0 S fd -p C D> to in P P 0 — t 0 P 0 U fd 4Z 0 0 ax: fd to u x» tn u p 3 -O > -P E rH u TJ cn a E C OOP —4 0) in a n E 3 X) £ in XJ 3 x^ \j m -r^ 0 tn C CP P c fO 0 0 0 C ■H q 0 0) < OJ C OJ a 0 0 -H x: 0 0 xj x: 0 0 P P OJ 0 3 ■H 0 C 0 >, c to in H 44 fd P -H to p fd 3 01 QJ 44 a U 3J<'Oi3--'T) ■H 3 to 0 u P P iw tM to 4-t +J u-i 0 in c E P 3 td tn w ■p 3 C QJ t4 to E cw*-* 0 0 43 0 E p w 0 0 CJ* 1 0 01 t3 -H c to 1 r- cr (U •!-> 0 •H i td 4J x: -p > ■H 0 rH 0 u m t3 01 C -P rd C at (1) 4-> W 0 tH Ti in (U -H c cp (d c tn 0 3 x: ■P 0 M ^ ■H 4-) ■H C n3 CO 0 >irH fd C Vi rl c •H XI 0 C ■rl 3 o >4-l j:: 0) U CQ -a IH rH <4H CO H Q 0 TJ ■•-' fd 4-1 4J 0 rH 1 OJ to 0 Qj to . ,U fd 0 u Cl tJ ■H ^j 4-t 0) IH U IH >i -p E tJ -P ■rl 3 0) rd 0) tn )H E tT"^ *^ 4J g m X -H rH nj C H 3 TJ 0 > rd 6 ■== fd (d -o x: C r-i at QJ M C -P rd ■rl s ■iH 0) 4J 0 C -P cn TJ 3 to 0 +j jj 4-1 0 ■H iH T3 0 -H C 3 0 4-> .3 <4^ i-i C M H to • (d td c -4 Ji^ 0 (U > OJ M TJ 0 .-H in 0 -P 0 C QJ ^ -P C tn C QJ 0 0 a TJ rH 'O ■H UH ■H *H > « Q) -H u -rl iCd -H -rl TJ rd to rd 01 -rl C -H 0) -H 4J X c ■H >i-U Uh 4-> ^ M -P > 4J -O at 0 3 C TJ 0 ^ E-^ Q) (U U to Ul (0 •H U 171 u u a u c Q) C TJ *j 4J 01 IH T3 U >i -rt CO LO tH • 4J Dj M O -P TJ nj C 0) 0 Q) ■H U • QJ nj ■rl M 0 01 QJ QJ IH ro C td M QJ c tn 0 -P ■P iH C ■P (d V4 cp w > ■ >. (0 c 4J nj O Uh MH > cx CO rH TJ C >H C Q) 3 C "w C rd ■H C -P M -H 0 0 < (0 o M U iH 0 0 ■H (d ■H C c Q) QJ 0 x: 0 Ot TJ ■rl rH 0) c at w tn 4J 0 (0 0 tl rH x: cn iH -H 3 at x: H o JJ u 0 ■H Tt *u T3 Q) x: -p u ■P 0 u UH 4-1 ■H CO 3 TJ u (d c ^ T) "tH QJ (0 QJ x: -p rd c c ■rl CP rd ^ U C (d Q c at (N X +J (d a 0 C 0 TI cu ffl to JJ -H iH 01 0 3 QJ -a a a u >i 0 x: . C -rl rfl a> QJ a .1 UH (d -O XI x: ■rl 3 QJ to E •H c u C 01 a 0) T3 T! >1 4-1 -H 4-1 < ■iH 13 g ■H V4 CO 1 0 tp +J CP (d (d UH x: E to rd Q 3 x: tH u ■rl 4J c 0 0 C to £ c tn tH 0 tJl ■p ■p 0 to 4-1 0) to 4J TJ at td u 0 C M (0 <1) 0 ^ CL rH ^ -H ■H C (d M c QJ c U QJ C rH to 01 x: 4J (d at ■H > 0) a. ^ j:: 0 x: (d C C C -P 0) u W en iH 0 x: CO Q] to 0 ja i-H x: !T> cp cp^p C a (0 W Oi c u (0 1.1 +j 0) c -H CO OJ 4-1 0) rH 3 c 1 rH 0 ■rl u e- C C C QJ rd in c c < •H 0) 13 Q. -H +J e H 0) XI CO rH C +J (d 0 rd CP to -rl -rl tH C e U C cp 0 o H 1-1 s C CU tji >- > (d (0 -H +J 0 H ^1 4-1 C Ot tn 3 3 ■P -H G ■rl (0 ■rl a c 0) 0 u (L> 0 01 J-> M W 0) CO q 0 rtJ - ■P +J (d C -H x: 0 (d . to 0 at rH rH rH 4-J Ul -H fl O 0 TJ ■p rH ja c td (d M Vi- ■rH tX E TJ C a 01 -p 4J a c VD 10 rH OJ T3 a rd TJ 0 d) ja M OJ u-i 0) 0 0) (d rH at +j TJ c > Q cr at Cl. 0 -P 3 4-1 4-1 i in 1-1 QJ If) 1-t oix: +J C - Q) (1) x: tj TJ •H c M -P c x: T) x: 4J QJ (d JJ 0 at QJ 0 •H 4-1 0) CO x: QJ QJ 0 4-> 0 - Xi n] U (0 0 x: 0) rH ■H rHl 4J ■n n3 0 x: > CT iH x: fd t/i ■p tn rd 4J ■H 4J 4J g > c +J .H c H x: -iJ U TJ -H +J (dl J-" X] C rH +j 0 0) 3 C 4J >.M 01 0 TJ UH C D (0 •H to G4 0 +J nj C C IH a (|H IH 3 1) Q) QJ y^c TJ 0 ID 4-> bJ -P IH UH a c 0 T3 (1) c 0 0 ■rt 3 fO -^ 3 ■^1 0 fd CO QJ n x: UH «4H ro -rl fO Q. 0 ■H 0) B^ **-! m TJ s-o T3 ojI a XI ■P 0 C rd tn to 0 rH 4J x: ^ s s 0 c +J (U (0 CO 0) C ' C 0 M c 0 u ■P c (0 -H UH 4J cn c c cn 0 Tt u (d 13 C ■H ■H 0 TJ CP 0 to (d u x: H c (d to 0 0 c C Ui c 0 H 4J XI rd IH 0 ■H -H ■H (d nj C >i b 0 -fH 0) m TJ 01 x: a u H ■rl M (d •H 4J 0 01 0) -H X C -H M TJ 0 ■H 4-1 cn 0.1-H i-i U-l l-i rd rH 01 ■P (D to 4J +J (1) OJ x: ■P •H jh xa E ■P QJ to Q QJ 4J 4-1 tn rH tn c <0 tji fl3 ^ -P I^ t-H 's- (d .-t ■H ^ to QJ C E rd u E CO Ti 0 (d ■rH rH 3 QJ Q S^ ^ p tu <0 bJ •H 3 to )H )H 0 (d 0) U QJ >i 1 -P to TJ rd 0 -rl at 1 0 QJ 3 rd u 3 u U 0) (fl U 13 M U 0 (d OJ m +J Q) tP-H u 3 ■rl 0 U -P 4-> 0 ax: --i UH ■rl tn XI S^ H (0 0 C x: c x: • to a-H QJ 0 fd 0 10 0 O. U ■P u rH M ■H CO c 4J TJ (0 ■H tj> CO i; a 0 -p w aTJ nm-t Q> > tM G. to C 01 UJ tP rd QJ C QJ XI 0 -o tn E-< 0) (Q (1) C > x: C to u g -H (d . U -P iH > ro c x: 01 x: -H c ■H ox: H Q) .C 0 x: 0 0 x: 3 x: UH *d x: < u CT (« (d H ■rl rd (d E fd u JH -P 4J ■rl (d a U -P w c u 4J jj +J U +J a H 0 < x: Eh 2 ee: < W 0 w fo H [iH 0 < X 0 > H 0 H Q H E- !- « 2 (a a. H U in Q) 0 13 i3 C X >i M OJ M >* 0 CO 0) i> u U C 0 E 1 XI CN 1 0) -H 0 0 P XI CO (0 -H nj .H 4J OJ 0 -H C £i Eh -t-" u p cn XJ -P c p c § 0) -P e T3 XJ C 01 ■H JJ u td ■P U 3 C CO • 0 CO a c iH OJ tp 01 C 0) 0 td OJ (d CO p a 01 P C d) nJ tn . -H x: ax: 0 ag 0 tU fC {X o P -rH C CO (0 rd 0) P ja -rH 4-> P P 0 c H 0 c x: M 0 OJ " 0 -P jj CO a -H £ 3 u P -r^ >1 p x: 3 w OJ 0 -P • ■>-* , 0 w X: P) M -H MH W 4J a a OJ cn 4J C nJ 0) (U WOO o. (d P OJ CO -H (0 0 P OJ -P T3 OJ ■P OHO • td X e 0 P 01 C rH C CJ -H Xi rH fd c P Q) (4-1 1-1 1^ P (c e u C U C -H C M n c oJ ■rH 4-1 C • XI ° 5 (U +J t3 c (0 u 0 0) (d c OJ jj OJ 0) C -rH W D T3 u-i 0 OJ iH 1-1 M — C 0 g fO tn Of Tl D TJ m iH c T) a-H CO 1-1 u E 3 tn 73 O) T3 0) 4^ <0 0 H in vh 0) 0 OJ c 01 01 2 in x: E M C C .H C -H o 0 T) 4J x: 01 OJ t3 C c (d fd CO x: CO x: iH ■H OJ ol -a rH OJ ax) a.x: p OJ c td QJ P cn JJ cn M O -u nj td a c OJ P U OJ c trip QJ E -H M 0 u <4-l 01 fd -H c 0 --1 x: .H 01 tj a » f-i 0) en C o u-i --H -p c xj x: c [i. M in 3 ■H ■rH at c o z > 01 0 • CO C rH -D u c: (D Ed C -H -P C-rH 1-1 iM (d Id td c 0 H 0 P P l-i 0) 1-1 CD x: -rH CO 1 x: P in CO •OOP Qj'O M-i ■>-i IT QJ tj- tU -H c 3 Cr>< TJ 0) td > M ■H 0 3 x: xJ P u p ■H QJ OJ c fd 0 to Tl *o CO X) x: m -H tn (0 OJ (0 c x: tp td td CO a P to c in XI U 0 01 c a- --1 u ■H QJ C M 3 +J -H D> QJ ^H 01 td td 4J (0 0 in o tp at <: tn ■H " CO tn 0 — C CT" a 1-1 1-1 (d -H td ^ -H > -H c U x: fO P c a !_, p • +J c Xl ■r^ cn I3^P C M C -H 0 C (U 0 fd U 3 0) M o (d +j ^ >,u-i 4-1 in 0 u &. c = ■i-i 0 E OJ C td 0 0) 0 P 0) ■■-t -rH xi x: P U -P c iH C 1-1 0 ■H tn 4J rH U-l 0 fd 01 -rH en 0) P to XJ OJ 4J E '"^ •H b. -rH «4-i o td > 4J n^ o (C ■H i*-i O (0 -H T3 CO 0) (d XI OJ ■■H 13 J3 P ■rH OJ E C C 3 J= 0 P OJ ^ ■rH U 1-1 O M V4 U-l (0 H c 0) CO OJ CO ■H T3 c 3 P >. to X) x: 0) 01 0 -H H trU -rH 0) U M OJ 0) ■P (0 (0 -P fl 0 •H V^ P ■rH e +J -H X: -iH +J fd C 13 ty 0) E en 01 0 to x; C P (U e -rH o C QJ rH cn (0 x: H CD U 0) e P D (d If tn c x: -P .-1 >4-l OJ 1-1 C CJ) c •rH at in c - p C --H (I) cn c c 0) ty E-* 3 0 CO (d x) -Q 01 -p 0 x: "4-1 cp 0 H •H • P x: 0 OJ 0 -H CO 0) M <0 tw 1-1 OJ (d OJ 01 iH 0) u< to c 0 U 0) ■H x; H4 P 0 < Vh > c td H 'H c u fd M x: > 0) i-i 0 3 1-1 ■H x: Q) j: 3 01 H C u -H X u 01 t^ c c ■H 0) ' 0 rH o -H ti U 0 -rH +J (D -P U-l U +J P +J in ij • 0 IW u l-l m >ix: 0 -H 0 a rH > ^H X) -H OJ ^H OJ at •H P iH 13 . OJ 4J fd -H OJ tn 4J 0) C C -rH U (d (d CO p P 0) O Vh -H x: a (u -a w p P o O X) 1-1 P e 0 p -H ■P -H a 3 0 XI 1-1 iw a u OJ 01 3 -rH O 4-1 -H 1-1 iH ■•-* c (D 3 ■rt 0 -U fc. P 0 -P 0) n] c ■P 3 ■H >iP td trx: -rH P ^ CO X) -rH (0 P 0 Cr 01 XJ -H P 14-) 0) ■rH 0 m m 0) -H 0) (0 C U TJ M P (0 u .-1 Ti cr- 1-1 OJ 4-> xi td fX CO c p c u CU tn 01 § a td u in at 3 u P -H 0) Tt 0) ax: -H U fH 1-1 OJ c Q -d Q c e g -H (d Xl 01 -r^ 0 0 to 1^ M V4 P r-l 3 > M c 0 x: Di ap 0) u -rH C CO <0 ■P 0 U "w fd 01 in ■M a-H 01 Id 0 C 0) fd 0 Qjp >ix; tp 0 e n & (d P C XI U-i 01 c *M a > OJ QiP o at U) (y 4-J P U ■rH U Vh XI xt JJ -H 0 13 in 3 »-i C E 3 3 0 C ■H p C H x: •ri 01 td ■H x: 0 1 a CO c x: OJ oi c c c 0-H CP C C ecu C 0 -H m a-H -H rO 0) WOP M r CO -H Tt +j U. 1-1 u .-H Eh 4-) 1 (d 0) -rH 4-> d; = M < -H ap fd fd fO<:D a-HXi-H 4-1 at p in E o CP OJ 5 Id c 5 cp a-**-! ■f-\ TJ .§ 0 c td OJ c x: rH 4J ■H 3 c CP m 0 td OJ s, 0 c P E X c a c o OJ o at 0 (d •-* x: •H OJ p P c 0 > to rd u 0) c OJ •H XJ ■H 3 > CT § C J= X) 0) p 01 at p ■rH XJ x: in »4H 3 XJ p >i 0 (d in x: 0 to u c p o c ot OJ •H 0) IH 4-1 OJ 3 OJ fd XI o o ■H (0 ■H c U cn rH P 0 O fd Id U u cn x: 01 M tn T3 Id 0) Id tn P 1-1 OJ 1 C 3 ' I rt P ' ■H 4J t-H > x: Id -H P C-H 4J 0 P o x: c fd tji tn 01 3 P P O 0 U O P • ^ to a p a E »p ; •H 0 1 c td ox: CO C c o O td 0) . a a Q y tn C 01 0 o OJ P ■ -H XI fd >x) P u u in cpx) OJ Id -rH C cn x: E to at 3 CO Jj O -H fd to -H P p at p u m -rH c l-l u at fd OJ -H > P X: rH c c I O O I ao 0) m X) OJ CP o 3 J:: C rH 4-1 -H C 3 fd 0 X3 > P C rH !h 3 Id at XJ 0 C 10 OJ ia 0 c XI ■■HO C OJ P u Id 1-1 fd Gi Id c 0) Su x: in OJ p c P c o c u 0) -H M O 01 P M-i I Id 01 c x: tn p o Id ^4 x: at vh p XI 0) c ail?: . e 3 c C C a OJ o o o x: ■H rH JJ ■ 4J jj 01 tn u td > XJ Id ot x: OJ c OJ tn P XJ td cn o t. in a -H CO XJ E OJ at E iH in o o u in a ■rH O 0) j= H x: E-" ap I I OJ 1-4 01 P U O ' o in fd V4 CU o w ■-^ a P CU OJ 3 o o > u 01 a I o ^1 -H -H x: in ^4 P J 01 u H P H 1-4 ■ 4 c Id Id H -H 4J 3 > C M J *w < X3 o c ■H Xl -H JJ OJ 13 at tL, t otCidUHodotfd c ap p p p x: . c P 01 o in 1-1 u ot . o ot P U XI -H ■ id3oicao^rHPui 4J -H XJ in in 0) x: U ^ C C 0) X) P CPX) OJ ^ M OJ C r .S^-; fd Id a 0) _ . . - . . Ei3 — toxjx:o x:atatr-4s rHO^HOt-HPCOC' >lp p at o OJ ^ Id rH c u ot 0) E tn tn r-i cn ■H Id - 1-1 G 0 OJ u x: -H JJ ij td - ap 0) c x: td-H0)lHfdUCOaJ'HU-.Q)4J P 4J0JO OOtOE aoioi'-H-'Hinu> c >• fd cnxJ^wPC 3 OtP 0X3 i:aj ojthooqx:^ CP-rl XJ u x: ot ^H to -H I •H > iH iH • rH c Id > o • O m u cpr- C iH C C-H— ->TrH OJ « Ccn-HjJ O M-H^lJMrH4JJJ^H-H.-H U-|i*j Ut- 0)td rHcnid tdiOP -H-uijP IdC iJlH CniHrHU3fOQ3 cO'-HOJotrH Id •-{•-* a ■r1 -H Id P X) Id tn tn iH a-rH M _ . . ■H OJ Id CO 0) P ■H H 01 > X) X{ C X) x: OJ OJ u o 01 xl C 01 a 01 ■rH xt td cn P at tM 01 p o 0 x: OJ 0 ex: x: 01 p ■H ■H u >; P tn cn at u c P ■H 4-1 iH p 0 0 01 -H X) 0) C P c Id Id td 0 c x: x: to OJ p td 3 ID c E S P 0 P H P tn — tJ ^j cn OJ o M j:: -H WPP r-t OJ W Q} OJ c Vj o -T) M tn «4H > 0 (11 -H 0) j:: nj w Q) ID O 0 ■'^ -rA (/] Ul +J 4-1 .H C o e x: cu • tn at 4-1 4-1 ro M tl,-H fd -H IH -P +j at £0 04-ioojto(iix: 'U u w T) 4-( x: • ^^a3oj-^x:4Jajoj>i C U 0^4-1 twtiXl -HrH -H TJ (d tH m 4-j r- OJ Q m ■H0)CC(dl-IO-'HOOJH(0 M C c > C (0 u tP 0 c xlOOtn 4-1 >4H <^ u X (0 ■H at -H IH C -H E OJ C 0 C 4J U -H -r4 >i . W 01 a Ci^ t-i 0 t( 4-1 >, 0 x: •H C-H 0 4JTJ'tnrHotntn'a c >; C QJ U M U -P +J O -P ■Hcoja ojAiM-P 'OJai 0 (U (1) >, O -P -U M n3 W -H U 4-iox:ooJrHtn(TJC3U[j^ ^ ^ ■r^ c c (0 3 0) 0 0) -P OJ UC1.4JX14-lja-iHrHTH OJ CT-C 4J XJ 4J OJ -H 4J i-( x: -p p» (d m 0)3 pfOnJMP <-HCTl c (y (DUE C ja -p tn 'oto4-)-P uojrH 0) -H at c t-t (13 CLU-I < at c nS -ri C 4-iC t«Q,lH-HE(04Jtn ■U J3 -H 01 0 o &4 -P 0 x: 4-1 -H 'DCnjul 0)0)4-10 OO m T) -H 1-1 01 OJ -H -H 0JO)ap'4HU4JtHuaJi::a tn tl > c U - 0) -P C -PTt tnTlXMOiJ(d(0 > 0 E5 tl) rd E <-t 0 TJ OJ ■HCQJO (OOJCLiOOltOti jJ C 01 'tn3tifd x: nj >i OrHOj-Prj 04-t IM • >^ 1J 0 -P u c to in ■H (0 TJ 0 < 0 TJ M W nj t( - ■P c tn.-H .UrHOl OJ>lS 0 M 01 iw V-i O U tJ l-i fO C -H tnCrHWtdnJOC >i<-H 4J tj> (A H 0) W -H -P 01 ot -P o OlOO] -CX+JtiO rH C ■P > c 4J 0 tn u-i c a-P ■H -rH C 4J -H U ID OJ 0 -H >, 3 C ■r4 U 0) nj Dj -P 01 ■H Q. c E +J Ot ■H (0 at (04-1 ••H Q4J014-JCi4-(O4-) fO C Q> M O ■-< 01 E 0 u o sz -U tj QJ Ul au-t 01 c a TJ -HO a o x» -D U rH -H XI oo-Hx;ajtoooE04-»oj .§•:! p. a « -P cr>r-i c ■p axi u H x: -H XI 4-i HrH tn H (U OJ 01 0 OJ m 0 tn jHdOl +J C3CU OU-i u 13 oa)jHi-(j-ij::-rHjn 0 trtJ 0 tn a ot 0 tn j<; >iXi oj iw 0) x: u x: 0 tn jj 01 x: o m.-c-H fOj-iaiH 0 « c -P c y-( ra tn -H >i c CO) tnotO-PC4-)--Htn >i -H 0 0) > U-l ■P 01 -Hx:-0JE aro rH Q) 4JC)H-H)i:ox:o-HOE to Op4JOXt4J4-'Xl-P-rl . C C u m -H W C u at 0 ■H C E 0 -H m W 0 W C 0 1^ u •H tl M -HO D OJ -H (1) tr MtM-HtO -rip (0 CPOt (0 Ul D U C > -H W 0 tn c ja s 01 [i]O4-)r00)4-'in4-'ttt]::T30JrH w c U 0 01 -P c tn -H 0 fO 4-1 Jh uoJE'^ cv4-rijHi^xi P O TJ u iH -rH ^( -H 0 OJ jC M 0) rHHfdE'oO4-i010J n] (D U ■r^ 0) fO nj -P in -H 1-j E-* -P -H J3 TD (0 WXl(0E:*'Cn'04-i m jj ■U c c cu 0) 0 tn C tn QJ ciHrHg px:e shoj a •HQJ(0-rlO)tn4JOOJOJMtiOJ ■H (TJ w ■H ■H -H ,C Qj tn tp -H flj'O tn T) -H o Cl. M H P C . 0 TI O 0 [i44JiHMx: ihe> ^ ^ QjOi aj4J4-t oi-ri-ri ojxi'a u 3 Cr w o • u ■H CO £1 ■H Q. r- OJ Jh 0 aj(0Tt4J f03>tj>S4JCnJ x: 0 p j:: x: , H td -H ^ > tn (0 3 4-" 4J 0) E O (TJ QJ o c tn p u x: > tn-H - frx: 4-1 fO OJ 4-1 u yn x: c TJ i:; Q) -H o S at OJ > x: Q)TJ0t4-»E'-<3 - ■H rH XI U +3 O TJ > p OJ Jh tn (/) rH O O rH (TJ QJ 01 3 ojx:4-»i4javiuo x: tn QJ OJ us 4J QJ Vi O 4-1 3 c E o o OJ J=o-ri>(Qjcms 4-> -H CT ■-< x: c 0) ■ri 4-1 OJ 01 4-1 nJ Ul S (TJ IH 4J U N rn tn iM tn tn -H c p -H c o iH QJ C O & 0 01 OJ n3 -H 0) 4J -H C 4-1 Jh Di4JTJi-t4-) O t CP M (TJ (TJ C -H 3 (0 O > QJ +J V4 w - > a '> 0) QJ 0) -H W C J3 ■ 4-" iH tn FH O -H ropc-PMUviw 4-' 4-1 O C 4-1 IH tJlrHU-rlrHCOlld 3 O TJ O -H 4-1 y-iUOiaC-rlTJC 0 -H x: -H 4-1 < Ul 4-' tn tj (TJ 0) 4J cr -H > x: E c: 4-1 O TJ (0 O 01 4-1 U Iji 01 aTl )H -rt (13 C 4-1 01 O rH U. (T) -H tn Q 0 t< 0) Q 4J 01 C^ O XI 4J TJ b. 0) ■(-! 0) (TJ rH ^1 x:2:'oajx:x:Qj(a hdeS4-14-iihx: Ul TJ 3 ^ O 3 OJ 4-1 C -H Ul 4-> tn O (TJ OJ ■r-ij:; tj, (TJ 4-1 (TJ E J-t tn M (TJ --4 O to x: Ul (d tn QJ XI ■H 4-1 O C QJ 4-" 4-1 (TJ C TJ 3 O rH tn 4-1 u 0) »44 01 tl (0 TJ ja a Ul c E 0) x: c (0 Ul 4J -H ■H E tp tn lui -H c OJ O -P -H TJ > C 0) r-t 0) ox; o x: -H &H > ■P -P c TJ O • c -H 4-1 tn (t3 rH C C 4-1 XI QJ O to 3 E -H Ul a E tn OJ o tn TJ M O p C 01 u p -ri >i tn f-H rH -rl W M 01 TJ rH (0 E (TJ OJ -rl U •H O 01 (TJ Uh i|H Ul E »4H 0 0 O 4-1 E -H c a rH OJ S 'H (TJ E O TJ ■rl -H 01 4J OJ 0 QJ 01 01 M x: 4-1 tn QJ 4J E (0 4-1 Ul a 0 Cu Ul 3 OHO 0) 1= H 0 x: cu-D Q -H W O .§■" (0 (TJ C 3 X: (T) u a a c a o ■rl 'O tn c tn (TJ P o c T3 4-) tn (TJ 0) 3 Ct- E t|H O -rl a4J 1= •-{ ■H -H 01 (TJ Ul x: 4-1 4ii 4J tJH tn >. 01 P T) 4J C C (0 -H U 4J tl tn 0) tn OJ u QJ c: U IH p TJ I4H i:: flj o 01 IT- a (TJ c < tl (TJ 01 x: U +J OJ >. c (T3 OJ 3 OJ x: XI 0) E o o 3 -H C 4-1 4J 0 tn u U >i OJ tn to 01 o x: o »H Eh 01 O Q) • tl T3 <0 01 (d 01 a to rH 5 c U fN OJ > 01 OJ 3 X: Ul -r-, 4-1 3 O -rl o 4J (^ r-{ -H -rl tl d 4J (d 0) tn ro x: in t3 TJ OJ 01 • tl a tl tn (d 0) -H arH 01 4-1 C (d w (d c 01 0) OJ -rl rH tn >4H XI aj3 o (d -H TJ tl u tn Ul >1 u -H in (d tl ■p 0 a 0 E u a 0 4-1 o u 01 „ (d QJ . . *tH 4-1 4-1 C (TJ >i CT>< U XI C -H ■H tt TJ tn TJ OJ C rH (0 x: -H i ■rl -H (d 4-1 4J 4-1 01 tJ 0 U W 01 01 QJ QJ Ul W W Ul 4-1 c tn 0 OJ rH •H O (d 4J M -rH C 3 U 01 0 M E « at 01 OJ u Tl a o tl a 4-1 (TJ (d 01 >. c T3 rH C rH M (d fO -H • a-H in X *J 11 a 01 C IH 3 01 01 O c 4J x: tl QJ O 4-1 tp OJ a XI -P 0) tn c tn to (d OJ 0) (d to j:: x; tn OJ 4-1 • 4J tl M c a c • (d -rt 0 01 x: 0) •H C IT tl 4J -H 3 0) U tl O 4-1 OJ (d x: c to E 4J ■H ^ s ■H M -U OJ +J a rt x: fo S ui -P rH 3 • CD a w C -P T) 0 +J 0 ^ 3 a 0) ■H IT] B > HO. > X 0) 01 01 {T ja o o -p W +J (0 >i c ■tl (N fO O < (0 a ■ O 01 H 13 > rH OJ O +J fd ■rt c -P V i 0) o ^ e H C C fO 0 W 0 O )-< -H ■H -H (U (D .-H p p-O > (13 to U -H 0) c o; (d w T) ■HP c Cl. tT"-! O J= (0 U P 01 01 tl -H ; X: £ 0)13 s I £-• p x) at I 0) w P I tP Uj -H C I • C > 0) I 73 ■'H 13 Q) P I -H T3 01 U to I i-< 1-1 tn -H I 03 to O C W ; > tj- a 0) c OJ O 0) 0 I 10 Pi t-i J3 o ; c a c I -H W -H d] • 01 tJ ■ e c X j= p S o -P o P -H 0) c w - p I D W O W u o Ot C -H rH ' ■-I 0 +J rH W ja u fo -H M tn u 3 w ■H tn o jj -H M 3 .-t 0 x: rH o 11 CPP (C >i c 01 -H c w i-H tn [lh I P lO n3 cue 01 O Q) O ^ SZ C --H -H U -P m OJ (0 x: c •H M c 3 -H 0) p ^ C XI C -H C ip -H tn x: C C <4-l P c ■rH ■H 0 ftJ 0 01 P CP -H 0 0 at to tn c; XJ en >i-H 0 3 P (D ^ tn to P ■P C C C P to u 0 tn H c «p to P 0 C >P O to to -P 0 1-1 0) X 0 to c at c 0) a ot -P > P c a 0) J3 tn -rH •H x; ■H M E at p OJ m 3 x: C M P T) ■»-' iw (0 01 g x: to e cr OJ .p • OPPt-HCiDPtO p C 0 OJ to c x: p x: a) T p ot a-P 1^ to -P p u M cn-p -P 0 • c 0) w a to O 0 x: c 0 3 ot p 3 ao -H XI QJ (0 a Q) XI U-i +J 0 ip p p to p at ex x: i-t x: Qj u ■H 0 at c 01 to 0 a> p to O M 0 -H C c tn -H p 0 0) E x: ■H to at au-i D 0 0 rH c rH at u x: x: at -p 3 s: x: ui 0) IW -H U 0 p n_i p OJ p x: 4J M 0) Ja 0) P T) at -H (0 01 C W -H -P 0 w p u x: to -H a> Tt -M OJ P 0 C 01 3 -P c c at +J -D 3 tn at to ox: Ti c 0 0) 0) .-H -H 0 T) ■ -H C 0 tn P w ot -H ■r4 in (0 c 0) T3 a c to P (0 in C71 C C 13 4-t C -H x: 01 ■H . 0 rH C 0 P 0 -H P to 01 -H > ' D tj' 0 < -P ■H (0 -H > 3 u u to t. c mow T3 at p P P ,C tn 0 &. 0 to 0 tji p • C 0 c aj -p p to p c rH tn at ot -H C Q-D (0 P in • at tn 3 tX'H • •H nH x: *J T3 ro to E 0 ot tJ 01 to x: ap* tn tn iH 6 01 at u 0) -p -H (0 ■p p c ot M x: M ^ to .H •- C -H at w p U XI Ui C -r^ • ■P T3 (0 P tT> <0 c to -H T3 E 2 C at -P aj OJ C -p p 0 p <1) g (0 u f-H P P P > (0 rtJ m X) Q to at Q 3 pp cx:s OT) ai-H c -H 0 -T3 a M at OJ u OP Uh C > P to U OJ C a ot u E Ot XI tn 0 ip C -H 0 to )-l > (0 x: j:: -h P 0 M 4H • -H CP > X 0 t/i -O 1 P 0 01 P 0 at P 0) iH 4J ^ T3 .§ c aj xa c x; x: 10 PC M 13 ai c OJ E 0 •rH (0 0 E-* 'P +J c C -rH tJl OJOPCWCPOW S n> ■H ^ -H 0 0 0 3 C "p-HOJO rOtOCC p Q) p P in p Qi •H ■p d) P P -H rH ot 0 w u W to m (0 • aj c ■P m p 0 ptoOjpratnhoiu >1 at at -p t/l U -rH (0 P c cr • 0) (0 to C -P ot -P in c & U --1 M C P c c at c at OJpXIP^PCPfOO) w&U'0[l.o)C3x; (0 a e -p & p 0 c aj 3 bj to (0 p 01 i 0 J to ■H rH 3 (A 3 a ot P 0) « c at tn ot ■p 1 U 01 CP 01 01 t/a OJ at g ot H-l rtJ x: a c 0 x: m p c o 0 x: ot rHxiM pX: oipTJ c 0 > tH 3 fO U J^ -ri cu ■■H U T) 4J P cu P w ap u 0 «3 o to 01 (0 0 p u p 3 a TJ CP 0) at c 0 x: ■p OJ 0 p p x: u 01 ■p a. c 0) ■ 0 E x: x: p u ■p u •H 3 c -p 3 tn ai rH XI tn jj 3 3 3 a 0 u at to T) >i P ■p P 0 TJ > P «p -H (0 .-{ a to a a P T3 0 0) O .H Q C 3 p 0 OJ a) o 3 p x: T) nj p 01 01 in 3 p c CP P 0 >i ot •p P at > p ■p T3 i to •p •p at 01 I E cj ^ rH +j . .p -H e to p u p 3 c (0 • P <0 0 O G4Q »p E OJ -H 01 to Q 4J Q - p 0 to C T] -H c c at o p o p x: p tn u at p cp- p • pi C *J to u C C -H c p • -H (0 x: OJ c Q tp to e < 01 ai to -P -ji: XI s p 01 C P (0 rH o 01 c a XI p 01 E -H 01 ■H CP P ■'-' Q tn c 0 cp cp to ■p uj 01 c ot 0 x: at p -P x: a to XI P p to at 01 01 S CO rH at c x: r- XI E 0 P C (^ ■H •H rH to (J en c to -P P O ot - O "H c p o axi 0) T3 CT* tn p C C P P rH 0) -H to 01 u > a xt c ■H o tn p •H tJlrH C S OJ O E OJ tn > -p P 3 01 p u to 01 73 to -H ■P -H H tn > c -p XI tP ■H O 3 c 01 cp a. ■p x: p ai P H C C 0) 01 p rH 01 0 d) Si E • u x: to to u P en 01 M to T) WW C OJ OJ O -H rH x: in p 3 P -H c c o x: ■p o c UH P -Hi; 0 c P IW OJ ■H - tn o X «i >i P (0 0 -H au -p arH (OP OJ C T3 o w C -P ■P to OJ to P PC to x: P o C U -H u 3 at • c P >i-H to p p p (0 (0 c to 3 O M C P U W O XI -H at at p p t^ u> c ot -H 10 to p ■ tn Q in in 01 (0 o 0) ai I x) p -p x: u p P P o • OJ (0 P w xj -P c a • c P o tD to O C U tPTJ 0 Ot 01 at -H x: +J c > P -D to E to c OJ B M 0 (0 -H x: w 4J ap in o H C C O Tt (0 0 0 01 c •H -H P ■P p p • in «-i p (0 tn OJ ja N p p aj ■p -p OJ &x: p C P 01 P P (0 P P to |j> to p ■H P g O O T3 o x: »p U 3 - Q -P ) £-P CU 01 C -H T) > P O -P -H -P (0 at at . . tj\ -H x: u u c I rH > r- (7* C I 0 -H o> OJ ai o i tn "H rH c E -P , a aj D at 4-1 < tn XI (0 at at c p c x: ^ (0 in ■H p p u P u- P f 01 01 c 0 3 c ■H o ■ o ■■ TJ .-H p p in OJ P (0 U rH -O IP > P to -H ■H to P > 13 P -P -O O C Q. C -H p « < > a >i tn C 13 01 rH 01 -H C XJ U P H (0 U T3 H 01 c at -H rH 01 pi rH -H OJ CP to fO 3 T3 (0 OJ to P o c E -H 0) -P O (OP -P P p c O P c «> -H tn p 0) T3 rH XI Bai in (0 -p &- ^ u U (0 P P (0 -O p c to a 0) T) OJ -H 0) > -H in -D Q O -H 01 "H 0 > T3 P -H x: c c 0) 3 H -H -H 3 &- ja p >i auH (0 rH o 0) c OJ at 3 p .!<: J.; w at -H to 01 - P rH P > (0 -H -H .« C P CO ■p o - 0 [1. -P in H c n] 13 P -p 'O p p (0 nl C O 0 P E iJ E 3 ■ >i (0 U P (0 (0 tn -a O OJ p OJ p H x: (iH x: at tx-P (0 p x: X p 4-1 at tp 010 0 I P at c to OJ x: w M x: c P P o ■p p p > (0 in o £ 0) p p > a p ot (B o p x: p 3 to 0) tn c 4-1 ■H 0) fO rH OJ o 3 - e 4J 01 13 (0 XI > (0 at p at c (0 tn ■H 13 tn 01 p 0) (0 0) P o P P rP CP 01 rC Cb (0 ai T3 o Sc -p 3 P I to to (dec o j:: 01 -H U Q- P (0 0 (0 E P rH to rH 0) -H ' • < OJ > 3 CP >i P P 0 -H 3 at "p p c 01 o ■ J rH -P XJ (0 p .c • J XI o P u J -H OJ to C - : tn a a at o c . H tn in OJ 0 -H o O Q C +J -H } a 01 at (0 H TJ 01 P P> ] tp p .C 01 P ) 0 -H 4J IP Ot e u 01 i/t P C 13 P C : in -H c o (0 u o O -H P (0 P in (0 >i ' o tn UH OJ ) OJ c c -H x: J > 0 U Eh •H Ti ■•-* tU < p ^ p a ) to p to in • J p 0 o in J Id 3 o j:: c J x: rH u o ) X P rH p -H 01 o m 13 at 01 p x: li^ c P at 13 g P aj cl 0 a OJ -o o > at P 01 01 axi c o a 4J -H a Id >i ■)J 3 4J 4-1 01 4J C M 1 u c 0 0 C •H IM 0) 01 C 0 01 Id C >i c c c •ri •rt ki 00 4J -H ? O -H M oi-rHco](d.H 0 x: 0 OJ 0 o • 0 -H n) 4J 3 4-1 E4J-<4 01t-HOU U ■H ja -H ^fn 0 a 4-1 > x: TJ 4J 4-) 14 CP Vj to o OlEidEOl Wi4-i -H cocCuOtd j::tMj:: 4J 4J 4-1 • 4-) C c •H 4J 01 1-1 c Tt 0) C (0 C 01 nj TJ fl < 0 •H c 01 O c 01 0 3 to 0) 0 0173 'doi4-)OJ D]i-tCj::(ooi 01 0 OJ C (0 -H 0) 3 ■H a 0) g 4J c 0 (0 f* 0 -H M -H ti U T) T) EJ^CTJOIMCU-HJ-HC 01 x; m 0 -H to 0) » u 01 x: 4-1 -H X: OJ 4J 01 0 c m a (0 C a a -H £ 0) c 4J u 04J>utocc(d 0 0 U 0 m a ^ 4J o c J-) 0 a 3 fl 0) 0) 3 ■ri ■H C 01 4.> 01 -H 0 -H J3 -O U -P -H -H 0) o 4J u <0 m u g £ ia c -H a 4Jthx: oih uj:: c •H rH §ti u a 0) c (U u 01 u tpu U 4->01> 01 4-)01ld 0) m Q) 4J (0 A •H il j:: O M TJ -H c c MC .H0)C-HrH.r4Ji > - c g,s tu M J^ (0 ■o TJ 4J «M O -H 10 -r4 CO «jOCXl-H04::'-HS4-iid •H C 0 0 O rO U Wl OJ c 3 OJ V4 4-1 -H -H -H j:: -r4 H -H 4J 4J 0 -H •D *J 4J -H o > •H 0 - C - 0T3 - 01 C4J (n04-> JOJCiQ 0 -H 4J C to 10 -o u s 4-1 01 O C U 01 0] rt:aioJO)(ou EOtI 01 cp 0 Id ■ • -H CJ -iJ 10 C -H 0) TD 0 c u j: 0 4-> 0) -r4 0 0) E 3 to m -H -^4 3 >H U C 0 ^ 0) • +J X) -H iQ 4J n C 1 OJcridO-'H t yirrCT3*HidV< •H tj^ (d ij fN 0) Ul s a> o)mx:oi tT'C.ct7^4J -h-he -ojoic 01 (U ^ C u a • ■o JJ 0 C o TJ M 0) 0) U U T) 0) 0 4J3w-P-'-;m r^ +J « -H ei < (ij 0 4J •H 0) (Q 0) •H U l-« U M k4 c 0) 0 :» lj 4J 4H (0 C iH 4J +j • to 0) > 4J 01 0) to 0 fO 3 « > T) roc aitoajoJ4Jid 0 tH -H 3 to Oi C t-t ■H M TD T) H c -H >> u tr ro 0) 0 TJ 01 coi-H en ^'jz s: -h 01 a CI .I2> > U c c 0] m c j:: T) (dx:x:oii.iO)mE-i4J^ti H 4-> -H 01 In ^01 C T3 «w aE a> 0 01 m c 0 0) 4J o Q) •H c 01 (y (0 4J c 0 01 0 4J 0) C 1^ u •H 0 V V4 C •H X JJ (0 U( Li JJ (d to 4-) tp t-i 4-> •rf b, (Q >M 0 > .H > u 0) 4-) 4J O n U4 Q) JJ+J+J-OC •O'-H-H 10 -O S 0) £ C7i»M -H c c 0) •H u 4J fl] C u O > 01 C CJ. 0 ■ U •<-H(Oai-HC'W-HM r^ C -H 4J OJ 0 M V4 vj u 0) 0) fl c 0) -H C (0 01 i-ioi atoji:o -Hu 4J 0) -U «) .H •H -O U . 0) J3 ia C c 01 4J C 3 T) > ^ < S-H n c m 0) *u 0) c M (d T^ (0 -H 01 -H O 01 in g a 01 8 0 M 10 •H 01 CO JS 0 a> 'O •H 01 0 U OJ O tj"~i 3 C 3 OlS3>-HJ^C-HEC*l-i ^ t4_iai4->ao4JV40-r( %4 M > 0 (0 ft i3 c nJ *t-( -lJ j:: J3 Ui to c J2 TJ 01 ■H -H tr 01 > moiSo -H00-H4J a a O 01 0] rriiM 01 C •3 01 oix:di3x: j::-ho) -hoi 0 -a 4-> V c m £ > j:: iH 4J ^ 0) a 1-1 O V4 0) 0 01 4Jid uHuccno-H £-0 0 ■H a 41 c a-d 0 ID nj > n X 01 -H (0 Id E 01 Xt -P 01 E ■'' 'i-' s 01 o M OJ C 0) 0 O H 3-H m -H TJ 0 oi-o < •w n CD j: u C CQ g w 0) (D 0 4-) C4J-rHC»w TStM-H-DtO *J rH 4J « S^ c 01 U 4J i! 0 •H m E U 4J tp«w 3 (d (d 0 4 01 ^ -H C 0 -H . 0 4J o u 3 CO (d > « > > > ■H >' 0 > 0 01 01Cn-HU4J0J OC0JT3 -COJ U C a CO (0 c M 4J IM x: a s e 4J -H L< 4JEC4J01C>ti>0> (0 a -H a to 0 S to^^ c a> c c CJi c m c: c c u 0 0) ^ 01 4J E 0) 01 2 -H Id a 01 -H 0) M -H OJ 0> 01 -H •^^ 0 Tl 0 0 c 0 1 0 •H 0 4J 1-1 -H 01 rd U 01 3 y 0)O4J-H C0E4J4J ni4->4-) UM04J 304J (dU rH l^ * C n3 -P •H •H -H -H ■rl •r^ ■H -H ifl a > x: 1-1 x: 1-1 j: E 4.J 01 0 Id 4-1 -H c rH S 0) 10 c ■H ^ 4J 01 4J 4J T) 4-> o *J (0 4J u (d oi-H>04Joioioi 4->> tT.flMtM£Olt(T^ c c c *x: triw 0) 01X1 0) ^ 0) t) ■'^ w U c u U 0) 0 «4 0 -H U OJ 0) T3 -H 4J X c 0) x: c 0 3 ^ rH fl] -p -a £ 0) i! V4 IQ 0 0 0) 0 0) 0) c IU£ 0) c s: £ m -H (1) 0 C t^ M < >; j= CO u U) W U4 W -H tn +J W-H e 4-> O Cm :» < 4-) E JH -HQjocjidOJidtooio > g C « . ? OJ 01 0 ^ I tn 0 -r4 0 01 Id -H U 00 J2 CP >i fO ■o 01 C C 4-> 1 m ^C IH H Id 01 4J 4n OJ C > -r4 01 -PC OJ c 01 -H O T3 -H O to >1 X> C -rl — OJ W 01 01 V4 -H iw > CD -H -H x: 0 CO to (Q a c 4J -H >irH 3 Id 0 x: Id -+J tr •H 01 • * U 01 ^ ^ rH 01 > 4-) P ■H 0 -H C 0 Id H x: 01 4-> - u u 01 -H 4J 01 IT>rH 4J c IH C^ T 3 4-1 0 O > oil 4J a -H U 4J rH (0 C 01 0 01 01 4J > IT C -H 0) -H 4J a 0 -ti. a 01 u xi si 0 e OJ • 0 Id 4-) -H -H c a T3 01 04 >i Q C -H Id i^ -H 4J 0 M M 0 *" < ' u Id c Q, C 4-1 01 ^ nJ 0 V4 W -H 0 ct3 g XI 0 3^ Id u o • ro -H to -H Id 0) x: atH 01 OJ>01>iH-H 4JrTii^lH m M OJ 4-) Q a 3 M 01 01 V4 C M 01 4J 01 ux:x:-HE 4-i>iidoi 0) H4-14-l-3P0rH3lH o a 0 M 3 TI <0 OJ OJ rH V^ > CO > x: , M cpx; ^ Id 01 U 01 OJ 4-> a E TJ OJ u (d 4J 4-1 -H 4J E 0 -H CJ Id P OJ ja x: 4J U tr OJ CP 01 4-> »moioio)(Dj<;-hc E 0 Dl C 01 c OJ td x: a 01 C C 4J (0 4J PI > (d 4-1 »4H 4-) ■rl 01 -rt 01 M Id 01 4J 0 •r4 flJW O C 3 >-H -H 4-» OJ c Id . O-nlH C-HOIOTJ 01 -H « E an rH 4-> 9 4-t O C M -H O 0 (d TJ Id •H 0 V UK ja S 1 rH -H c -o x: (0 x: M E k4 C c s a V4 u -p -H -H X: r-1 to (d Q 01 C O C 0> C 01 4J (d OJ 4J M 4J D M ^ 4-) -H u 0 iS 0 (d P u 01 u 4J > 01 -P 4J Id a <-* c i! Id 4-1 01 -H 0 0 -H 0 00 -H Id c U 01 u 01 00 01 C 4-> 4-> H 0 -HU-I (d V •HOJ-H>C-HX:> 4J x: 4J *i u m -H c x: CO i 01 OJ 0 ^ OJ 9 tH 0] 4J ^ C fl) •H rH 4J O t4 Id OJ (0 ■H DO (d m 4J c (d i 01 ^ g C A3 -H OJ CL4-) Id OJ 01 4-* 01 0 4J Id -H x: Id T) u -a 4-J » u 0 01 1^ c 4Jx: Q ^ 3 OJ 4J M ■H > x: -M -H c tn V^ t-t £h 4J M C •H xa 01 c S *J M 01 -H C to OJ -P rH g 0 H > x; M c 4-1 0 OJ-H ■H 01 0) TJ 01 0 s: U C 4J OJ 3 « (0 4-) 01 -H (0 u Id 01 E " 0 Id ■H ti D> 01 E c x: 01 •H 4J cr OJ 4-> ■H -a OJ-H 01 0) o c 1 *J u •H OJ 01 - 01 ttH 00 -a -H Id S Ul 01 **J 0 T3uox:v & c to o-Hx:'Oto 4J3idi»H 0 4J u OJ 3 •r4 TJ « 4J 4J 01 C 0 0 01 •Fi-'i C 3 0 -H x; -H 0 u to i^ 01 01 OJ T) "Id rH 0 Id TUm 01 0 c 4-1 x: c x:-i 0 3 HO 01 ■H 01 rH 0 -H m ^ 01 V4 M iH a >iTl «H • 01 01 (d E -H OJ 4-1 4J 01 4-1 OTJ U C to >i 0 TJ P M Id 01 «4H •H (0 CO 01 0 01 01 u« 01 a c (0 01 IH 4J OJ o S -H OJ OJ -H t4 4-t W Id a Id OJ >EQ>Ex:oJOi 01 a 01 4J >: > ux:o5'dOio-H Id 4J TJ 0 4J c 01 Ul 01 TJ C •H M c 0] 01 O > -P -H (d w O J3 01 01 IH C 0 crP 4J il o >i 0) (0 4J U iH 0 4J 4-> OJ-H CP 4-1-H M 0 01 01 o u a 3 01 Id ^01 01 OQ 01 o ; 4J »W •H p 01 to 01 x: Id 4J « E 01 +j -H -a C 0 <0 3 •iH 01 -H 1 ° jC Id M rH 01 OJ-H ■H 01 « a 1^ 0 >4H Id c IM »K OJ c Id O 01 01 T) T3 rH ) OJ 3 01 J 01 " x: •-* ■I-' C U C 4J U (d J3 -H -H rt w CO a? T> 01 CP ^ to (b M rH C 3 00 O 01 X: T3 4J -H XI •u CO OJ tH OJ 01 c u *J c V4 -H E Id 01 -H < C 01 M 01 Id a 01 -5 0 4J c +J 01 -so OJ 3 01 %4 01 x: 01 •H 4J Q 01 OiQ OJ x: 01 -H « < M > « 0 4-1 W TJ x: to £ 01 4J 4J E C T) JS ■H x: 4-1 01 4J n 01 0 to 01 rH ■H 4-> 0 - U^^U Ut-HUH OM'H C U 4J Ufi TJ OJ rH 01 +> m •-i 4-1 M n ex: (du-ioiinTJid 4-> c p >i OJ O+JQCWrHOO 4J Id (d c 0 OJ E "3 tJ +J iJ jj 3 c Id C 01 o +J • > o c •d 01 4-1 IH TJ Q. O Id -H -r-i OJ Id x: iH -H 4-1 OJ JJ OJ 0 il£ a CM (Q -H 01 0 c n 01 OJ C )H(0«i®+j>pid'Dx: 01 4H (d M C rH -H v^ c 01 0) C O 0 OJ 4J OJ X} -H M 01 rH ■H OJ fl O T) '^ *w Id E •H 4J 01 4J 0 C TI ■ u Id 01 u S* OJ 01 01 XI 10 01 -H -H q-( u Id 01 P OJ U J3 U O u rH ti 10 m 4-1 > 4-1 4-> 0 OJ 4J ■ri cue c m T) OJ a 0 c ? g 4-> -1 £ 14 4-* t« c u 0 Id 4J a Id 0 M -P CidiOTJUC-HOl O OJ *d 01 C -H C C 01 c to V£> 0 01 3 i4 CP4J 2 i-H 01^ to 3 10 ■HUi'O'HQ-HOX: M M rH •H 0 U C7> 01 c Q rH u o x:^ 01 OJ 01 rH to Id 01 0 01 OJ (d u •H 0 U OJ C 3 Id rH 4J a 3 3 •H -H -rl E •H a -H o a flj OrH •HTJ > 01 4J T) 01 OJ to E 01 aTJ 4J -H a 01 D.-rl JH CfWri 01 <-H us: x: 01 OH 0 c x: a 01 01 0 01 x: 3 c CO x: 3 U H flJ 0 4J ti iTj Id (d 4-> 0 13 Id E- M ■ U^t00]4JM01O 4J 01 U< t^ O H T3 O < 4J 01