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of trees has not been assessed. For example, a molecular tree of all living things6 produced by K.G. Field (Oregon State University, Corvallis) proved highly controversial since the tree turned out to be supported at a confidence level of only about 50%. Additional data presented at the meeting raised the confidence level to 57% or 81%, depending on the tree-making algorithms used. One technique for confidence estimation that is gaining ground is bootstrapping, a non-parametric resampling method, entertainingly explained by Norman Gilinsky (Virginia Polytechnic Institute and State University, Blacksburg). Random samples of the whole data matrix are extracted, and trees are reconstructed on the basis of these subsets of data. The resampling may be repeated many times 1000 times may be typical - and the
proportion of those bootstrapped trees that reproduces each branching point on the master tree may be noted. Values of over 95% are taken as representing an acceptable measure of confidence that any particular part of a tree, or a whole tree, is the most accurate tree available, based on the existing data. It became clear at the meeting that evolutionary biologists should base their hypothesis testing on independently derived phylogenetic trees. It is also clear that the theory and technology of tree making are advancing rapidly, and that all biologists should appreciate the possibilities and the problems. As Stephen J. Gould (Harvard University) noted in his plenary address, phylogenetic trees are representations of contingencies, or sets of historical accidents, but should not therefore be eschewed as unscien-
Conservation ofBiodiversity: A Southeast AsianPerspective Kamaliit S. Bawa, Richard Primack and David woodruff THE TROPICAL RAIN FORESTS of Southeast Asia are among the greatest reservoirs of terrestrial biodiversity. For example, a 50 ha plot in Pasoh Forest Reserve in Malaysia is known to contain 820 species of trees. Within one particular area of just 100 m by 100 m, there are 210 species represented among the 500 trees. These data were presented by K.M. Kochummen and his associates (Forest Research Institute, Malaysia) at the International Conference on Conservation of Tropical Biodiversity, which was held in Kuala Lumpur in June. The meeting was organized bythe Malayan Nature Society (MNS) and the Ministry of Science, Technology and Environment of the Government of Malaysia, and sponsored by, among others, the US National Science Foundation, to celebrate the 50th anniversary of MNS. The purpose of the meeting was to review the conservation status of the unique biological resources of the
Kamaljit Bawa is at the Dept of Biology, Universityof Massachusetts, Boston, MA02125, USA;
Richard Primack is at the Dept of Biology, Boston University, Boston, MA 02215, USA; and David Woodruff is at the Dept of Biology, University of California, San Diego, CA 92093, USA. 394
region, particularly in relation to development and sustainable management. Ironically, in the still heavily forested countries of Malaysia and Indonesia, it is often forest management practices that pose a major threat to biodiversity. Often, for every log harvested, dozens of other trees, small and large, are destroyed (Illar Muul of Integrated Conservation Research, USA). Overharvesting is another problem (see Box I). As D. Poore (Inverness-shire, UK) pointed out, a recent study by the International Timber Trade Organization (ITTO) suggests that in Sarawak a reduction in timber harvesting from the current 12 million m3 per year to 6 million m3 per year would be required for the forests to attain a level of sustained yield. The IlTO study could not identify any country in Southeast Asia that is sustainably managing its rain forest. According to Poore, the conditions for sustained management of tropical forests include longterm stability of logging concessions, control over logging practices, an appropriate financial environment to ensure reasonable profit, and adequate scientific information to establish management policies; yet these conditions exist for few, if any, 0
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tific. They are the keys to determining why life is as it is, so diverse in numbers and in activities. The crucial thing is to attempt to find which is the correct tree out of the many possible ones - the one that tells us what actually happened.
I thank Mark Wilkinson This work was suooorted
’
GW87912.
for comments. bv SERC Grant
.
References 1 Faith, D.P. (1980) Cladistics 5, 235-258 2 Archie, J.W. (1989) Syst. Zoo/. 38, 239-252 3 Archie, J.W. (1989) Syst. Zoo/. 38, 253-269 4 Archie, J.W. (1989) Evolution 43, 1796-1800 5 Sanderson, M.J. and Donoghue, M.J. (1989) Evolution 43, 1781-1795 6 Field, K.G. et al. (1988) Science 239, 748-753
sites. Therefore, it is not surprising that the complex forests of the region are being degraded into secondary forests or being converted into monocultures of rubber, oil palm and other forest and agricultural crops, with a concomitant decline in biodiversity. The squandering of biotic resources is not confined to forest products. Pisit na Patalung (Wildlife Fund Thailand, Bangkok) provided several examples from Thailand that epitomize the conflict between development and conservation, and among different branches of the government. Thailand earns approximately two million dollars annually from export of snakes, but spends many times more to control the rodent populations that are partially regulated by snake predators. Other speakers documented the decline in populations of beautiful montane rhododendrons, economically important rattans, vines, and the strange Rafflesia plants’ that bear the world’s largest flowers. The highlight of the meeting, however, was not the pessimism about the decline in biodiversity or the misuse of forest resources, but the cautious optimism expressed by many speakers about the future of conservation in the area. Although the natural forests are being logged or converted for other land uses at an alarming rate, a relatively large area is being reserved as state or national parks. In Thailand, where a complete ban on commercial logging was imposed Elsev~er Saence
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in 1989 (though this has resulted in heavy deforestation in the adjoining countries of Burma and Laos), efforts are under way to increase the protected area from 10% to 15% of the country (Pisit na Patalung). For Malaysia, Tho Yow Pong (Forest Research Institute Malaysia, Kuala Lumpur) suggested that a matrix of nature reserves should be delineated in production forests to conserve the biodiversity of a region. The idea is akin to the proposal for establishing a network of ‘genetic management units’ to conserve and manage gene pools of forest tree species in areas subject to heavy commercial exploitation in the western United States. rho’s idea, if followed, could further increase the amount of protected area. The protected areas must be carefully positioned to include the range of habitats necessary for the maintenance of wide-ranging species such as hornbills. One of the simplest stratsgies is to include an elevation gradiant, ideally from the coast to the mountain tops, such as at Gunung Palung in Indonesian Kalimantan. The role of special habitat types, in oarticular salt licks, in maintaining oopulations of many large animals, such as primates and elephants, was reported by Junaidi Payne (World aide Fund for Nature Malaysia, Sabah), since sodium seems to be a uey limiting nutrient in many tropical Forests with a high rainfall. The increasing use of remote sensng for conservation and managenent of natural resources is another oositive step. Kam Suan Peng (UniJersity of Science, Penang) described IOW Geographic Information Sys.:ems* can help integrate data from nultiple sources and spatial scales. She reported the progress made oy MNS in establishing computer ,Jatabases for ecosystems in order to nonitor change in land-use patterns. Through remote sensing, many ,:ountries (e.g. India, Thailand) have earned that their forested areas .3re much more restricted than indicated by ground surveys based on ‘:raditional methods. Consequently, *:here is pressure not only to conserve existing forests but also to bring additional areas under management. Clearly, Geographic Information Sys1:ems combined with remote sensing &arepowerful tools for conservation biologists. One of the main economic justifications for establishing national parks is their role in generating revenuesfrom tourism. The tremendous increase in visitors at Bako, Mulu and Mt Kinabalu National Parks in Malaysian Borneo illustrates the great
Peninsular Melaysia, which the 19th century coloniel administration linked to India in terms of its value to the British Empire, developed conservcltionl%gislationearly. The first game law, 8&&t r%strictedto c%rt%inbirds, w%%%nact%din 1894; the first sanctuary, the Chior Gam% R%8%rve,w%8 est%~~$~d in 19Q2. A turning point in the dsvelopment of conservation was the 2830 Wildlife Cemmi&on, which inwired into the confficts between wildfifa conaarration and th% 8xp8nding sgriculture sectw. ln p-wticubr, the lowland dipterocarp forests w%r%being clear%d to m8k8 w%y for ptant8tion crops, notably rubber. The re~mm%nd%ti~~8 of the- commisskn E%din ?938 to tfie first n%tionef park, Taman Negsra; this encumpsssed 434000 ha, across thrse states. Sar8w8k, w&h perhaps the richest flora and f&ma in all A&a, has now the most coherent conssnration plan but the small%& am8 under consenratian (se% T8bte). Conserv8tion ansGtm%nt lags far behind praposgls th%re(though this may 8o%nch88gsf. With a @rang tmd&tan ofcustomary rights and a suspicion of government t8nd ownelship d%ting from thar time of Rejeh Brooke, hunting within rclserves continues generally unchecked and logging incursions are frequent.
Forest resources (million hectares)
Total land am8 Permanent Forest Estates Productive reserves Protective reserves National parks and wildlife sanctuaries Stateland Forests
Peninsular Malaysia
Sabah
Sarawak
Whole of MaEaysia
13.16 4.75 2.85 1.90
7.37 3.35 3.00 0.35
12.33 4.64 3.24 1.40
32.85 12.74 9.09 3.85
0.59 0.94
0.49 0.93
0.25 4.58
1.33 5.45
The first forestry legislation, le8ding to forests reserved for timber production, was enacted in 1930; but 8 National FareszPolicy, which I%dto the establishmcsntof Permanent Forest Estates in all states, ~85 formulated only in 1978, foilowing the appointment of a National Forestry Council. Throughout Makzzy&%,Nationat end 9&t% P8rks 8r8 the only category tagally %xemptfrom logging and agric&ture. Within th%P%m%n%ntFar%stEstates, ‘productive’ forest re8%rv%s8re destined for recurmnt togging, white ‘protective’ forest reserv%sare maintained for the protection of watersheids.St8t%t%ndFor%%&8re scheduled for conversion to agriculture. Timber hervesting, 8 major source of davefopment capital, reached its psak in the mid-19708 in Peninsular Malaysia. Sab8h end Samwak in Borneo, whose inland landscape is hilly and rugged, have been rapidly iegg%d since th%late 1970s. In theory, logging in for%%treserv%snow fottuws 8 8elective lcrgging systam, 8tlowing 8 minimum harvesting diem%t%rof 45-60 cm, depending on Sp%ci%s.M%r@gementplans are based on 8nticipated felling cydes of 25-45 years according to forast and terrain, but the sustainability of this practice in the long term has not yet b%%nad%quatefysubstantiated. Some terrestrial and arboreal animals eventually return and even thrive following logging, but many true for%stspttcies c8nnm edapt to such brbitat rnod~~~8ti~s. Many %giphyterp, including some figs upon which many vertebr8&8 d%p%&, c%nb%%xp%ct%d to dN8pprsar within the first few harvesting cycles due to lo88 of their old tre%-crown habitat. Penistetnt problems in implement8tion of a generally taudebfe conservation policy are attributable to the state government’s reluctance to gazette con%%w%tian8r%asthet woufd be administered by the federal gavemment, and to the b&m%#t%bteundemtaffing in government forestry and wildlife agencies charged with administering present areas, which leads to a lack of enforcement. In forest re8%rv88managed fur timber, subcontracting practices smang concttgsionaires of&n invalv% th% sate end r%s8i%of io@ging licenses, which Jiminat%s inc%ntiv%sfor tong-terrmplanning. The lack of a firm 8@%ntHfcbasis for the fereeasts of timber %COnomkXlSy 8ttr8ctive s%t%ctiv% system of m8n%g%m%nt,8nd unc%&?+in growth, add to the probl%m of su%.t%in8bte rn8~~~%~t. Them is 8lso 8 lack of a consistent and comprehensive program for invotvimgrur8l populations in cen%8rvation managemtznt decisions. Among competing UsBaof lend, con8%W8tion r%m8in8 the last priority, alter exploit%tiue mining, agriculture and f%r%&ry use8 have b%%nr%j%cted.With Maleysia’s current population 8t 17.8 miltidn and ~~icje8~voring tar@%familtes, pressure an natural resour#s is Hketyto increa8%in spit%of rapid urbanization. Now%v%f, non~~~nm~t%i urg~i~8tjons and conc%m%oind+vidoab have provided much otthe cat%&@for focusing 8t&ntkn on ~~~w~~~ 8r;td~v~r~~t5l protection. The M&ay8n N%tur%Society, 8s on%of %%v%r%f vibmnt cit.&n%”~r~~~s d%dfc8t%dto the %nv~~offm%fft, is s%en,in this tight, 8s a critic& instrum%nt ef &%ng%. Oth%m in&&% the Environmentgt Pro&&ion Soci%ty of Mat8@%, tb8 fUl8#%@8nFriends of the Earth, the Pentlng Con8umers ~%~%t~~n and th% Worfd Wide Fund for Nature of Malaysia. The MaE8y%nN&urn So&ty, who%% PresZd%ntis akto the Dir%ctor General of the Forest R%8%%rch In%titute of M%lay&a (surety 8 firat), has over 3ooo m%&i%m - in chapt%psin all 8t%t%8%xc%pt(%~~i~ca~y) Sarawak - and is in a unique position to raise pubtic cancern and fa8t%r setion.
Universiti SainsMabysia, 111100 Pmang,Mabysia PateriMtori Arnold Arboretum, Harvard University, Cambridge, MA02138,USA
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potential for eco-tourism in Southeast Asia. However, tourism has its costs in terms of environmental damage and loss of biodiversity. In Thailand, increased tourism has caused severe water pollution and degradation of coral reefs and wetland communities. In the Annapurna region of Nepal, where people have coexisted with the mountain forests for centuries, the forests have been decimated in the last 20 years, in part because of the increased demand for fuel wood by trekking tourists. A management plan developed for the newly created Annapurna Conservation Area, described by Chandra Gurung (King Mahendra Trust for Nature Conservation, Kathmandu), is addressing this issue by collecting higher fees from tourists for use in replanting trees, creating rural employment, and developing alternative energy sources for heating bathing water. Participation of local communities is the key to successful conservation efforts. This view was echoed by several speakers. Yet no radically new ideas were presented. However, the extension of agroforestrysystems (G. Michon and H. de Foresta, ORSTOM/ SEAMEO-BIOTROP, Bogor), the creation of People Development Programs to place all college graduates into rural communities in Peace Corps-like units (Clive Marsh, Sabah
Foundation), and education at all levels (P.N. Avadhani, National University of Singapore) were emphasized. Another apparent dilemma was the lack of trained personnel to manage effectively the protected areas and to undertake necessary inventories of biodiversity. Paradoxically, on a global level, there seems to be an inverse relationship between the amount of biodiversity at a given place and the number of trained personnel available to document and manage biodiversity. The participants urged the national and international agencies to enhance their efforts in education and training in areas pertinent to conservation and management of biodiversity. A highlight of the meeting was the role that societies like MNS are playing in the growing conservation movement in Malaysia and elsewhere. The MNS has transformed itself from its origin in 1940 as a small gathering of European expatriates to its present membership of 3000 consisting primarily of Malaysians. While the leadership of the society is still drawn heavily from universities and research institutes, the membership base has expanded to include people of all professions and all strata of society. Increasing emphasis within the society is being placed on the education of children in the conser-
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vation of nature, and the development of published materials for use in schools, according to the Society President, Dato Salleh Mohd Nor. Conservation of biodiversity will ultimately require considerable regional and global cooperation, as was emphasized by the Prime Minister of Malaysia, Dr Mahathir, in his opening address. The loss of biotic resources is not confined to the developing world. As Dan Simberloff (Florida State University, USA) pointed out, the rate of deforestation in the ancient forests of the northwestern United States is now twice that of tropical rain forests. Thus, the continuing erosion of biodiversity is a problem common to all countries, and it affects everyone since new natural products are used throughout the world. The presence of the Prime Minister and other high government officials, and the fact that the meeting was co-sponsored by several international agencies, certainly indicates that the problem of biodiversity in Southeast Asia and elsewhere is beginning to receive the national and international attention that it deserves. References 1 Ghazally lsmail (1988) Trends Ecol. Evol. 3,318-317 2 Haslett, J.R. (1990) Trends Ecol. fvol. 5,214-218
EcologicalEconomics Jacqueline McGlade THE FASTEST-GROWING INVESTMENT AREA today is the biosphere. However, it is not clear whether this is the result of short-term, local disparities existing between the social value and the market prices of environmental goods and services, or of real sustainable development of global resources. What is worrying is that in current economic terms, resources simply do not run out. The Brundtland Report’ echoes this by concluding that a fiveto tenfold expansion of the world economy is needed to meet legitimate aspirations, even though current resource levels probably cannot support a fourfold increase. Unfortunately, sustainable development is currently based on a mixture of theoretical concepts and
practical considerations that has yet to result in coherent criteria for management and policy formulation. But agreeing that the philosophy of sustainability demands a new form of economics is one thing*; actually making the changes is quite another. The debate continued afresh at a recent conference, ‘The Ecological Economics of Sustainability: Making Local and Short-term Goals Consistent with Global and Longterm Goals’, at the World Bank in Washington DC, organized by the International Society of Ecological Economics. The conference focused on the interconnections uniting the world’s economic and ecological systems under three topics: an ecological economic world view; accounting, modelling and analysis; and institutional changes. Jacqueline McGlade is at the Arbeitsgruppe Sustainable management has two Theoretische Okologie, Forschungszentrum fundamental bases: the physical proJiilich, D-5170Jiilich, FRG. duction of resources, and micro- and 396
macroeconomic theory. In practice, these economic and ecological research activities have rarely impinged upon each other. The lack of a common theoretical foundation was brought home by Herman Daly (World Bank), who found that few macroeconomic texts had an index entry for the environment; in Daly’s words, there was no pre-visionary role for the environment to play in our current economic framework. For many, such as Bruce Hannon (University of Illinois), Malte Faber (University of Heidelberg) and John Proops (University of Keele), the cornerstone of ecological economics is some form of accounting system, usually based on the input-output (I-O) analysis of Leontief. Anne Mari Jansson (University of Stockholm) applied this philosophy to study the opening up of boundaries in the Baltic, and Alison Gilbert (Free University, Amsterdam) to the economies of Botswana. Some key theoretical contributions included an ecological utility theory (Bernard Patten, University of Georgia), and a demand-side cal-