United Nations Environment Programme … · United Nations Environment Programme WORKSHOP The Last Biotic Frontier: Towards a Census of Canopy Life Scientific Report Royal Belgian

United Nations Environment Programme

WORKSHOP

The Last Biotic Frontier: Towards a Census of Canopy Life

Scientific Report

Royal Belgian Institute of Natural Sciences Brussels, Belgium, 5-9 July 2005

Convened by:

Maurice Leponce and Yves Basset Royal Belgian Institute of Natural Sciences

Smithsonian Tropical Research Institute http://www.naturalsciences.be/cb/ants/meetings/esf_exploratory_workshop.htm

With co-funding from

http://www.naturalsciences.be/cb/ants/meetings/esf_exploratory_workshop.htm

United Nations Environment Programme

WORKSHOP

The Last Biotic Frontier: Towards a Census of Canopy Life

Appendix

Royal Belgian Institute of Natural Sciences Brussels, Belgium, 5-9 July 2005

Convened by:

Maurice Leponce and Yves Basset Royal Belgian Institute of Natural Sciences

Smithsonian Tropical Research Institute http://www.naturalsciences.be/cb/ants/meetings/esf_exploratory_workshop.htm

With co-funding from

http://www.naturalsciences.be/cb/ants/meetings/esf_exploratory_workshop.htm

IBISCA: Programme 2006-2010: Towards a Census of Tropical Rainforest Life

A scientific endeavour to survey the inhabitants of

the richest habitat on Earth, the tropical rainforest, and to understand their ecosystem function and value

Contacts: Relations with institutions and partnership: Olivier PASCAL, Pro-Natura International, 15, avenue de Ségur, 75007 Paris, France, Tel: +33 1 53 59 97 98, Fax: +33 1 53 59 94 46, Email: [email protected] Scientific programme: Yves BASSET, Smithsonian Tropical Research Institute, Apartado 0843-03092, Balboa, Ancon, Panama City, Republic of Panama. Tel: +507 212 82 33, Fax: +507 212 81 48, Email: [email protected] Technical information and research projects: Bruno CORBARA, LAPSCO-UMR CNRS 6024, Université Blaise Pascal, 34 avenue Carnot, 63037 Clermont-Ferrand Cedex, France. Tel: +33473406253, Fax: +33473406482, Email: [email protected] Data management: Maurice LEPONCE, Conservation Biology Section, Royal Belgian Institute of Natural Sciences, 29 rue Vautier, 1000 Brussels, Belgium. Tel: +32 26 27 43 58, Fax: +32 26 49 48 25, Email: [email protected] Specimen collection management: Héctor BARRIOS, Programa de Maestría en Entomología, Universidad de Panamá, Republic of Panama. Tel/Fax +507 264-5431, Email: [email protected] Media relations: Roger KITCHING, Australian School of Environmental Studies, Griffith University, Brisbane QLD 4111, Australia. Ph +61 7 3875 7491, Fax +61 7 3875 5014, Email: [email protected]

http://www.naturalsciences.be/cb/ants/projects/ibisca_main.htm

mailto:[email protected]






Workshop:

The Last Biotic Frontier: Towards a Census Of Canopy Life

With co-funding from ESF and GCP Brussels, Belgium, 5-9 July 2005

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TABLE OF CONTENTS

Executive summary ……………….……………………………………… 4 Rationale ……………………………………………………………….… 6 Background ………………..………………………………………………. 6 The IBISCA project in Panama ………………………………………....... 7 The IBISCA workshop in Brussels and the future of rainforest studies ….. 9 Why bother? – The applied significance of a project such as IBISCA …... 10 Who is IBISCA, why is it different and who supports it? …………......….. 11 Towards a Census of Tropical Rainforest Life: Schedule and Implementation ………………………………………………………… 12 Towards a Census of Tropical Rainforest Life: Budget explanation …. 14 Overview and cumulative budget 2006-2010 …………………………..…. 14 Module 1: Rounding up the IBISCA-Panama project ……...…………...... 15 Sub-project 1.1: IBISCA-Panama: processing of ecological variables .. 15 Sub-project 1.2: IBISCA-Panama: taxonomy ………………………….16 Sub-project 1.3: Improvement of the IBISCA database ………………. 17 Module 2: Dissemination of results ……………………………………….. 18 Sub-project 2.1: IBISCA web site ……………………….……………. 18 Sub-project 2.2: Dissemination of IBISCA results ………...…………. 19 Sub-project 2.3: Running costs for the IBISCA network ……………... 19 Module 3: Pilot studies ……………………………………………….….... 20 Sub-project 3.1: Organization of pilot studies: scientific meetings ….. 20 Sub-project 3.2: Maximize current canopy access for research …...….. 21 Sub-project 3.3: Improve canopy access …….…………………..……. 22 Sub-project 3.4: Maximize the value of biodiversity information …….. 23 Sub-project 3.5: Maximize the processing of biodiversity samples …... 24 Sub-project 3.6: Maximize surveys in different tropical habitats ……... 25 Sub-project 3.7: Maximize collaboration with biodiversity inventories..26 Sub-project 3.8: Relations biodiversity – ecosystem functions ………...27 Sub-project 3.9: Applied aspect of biodiversity research ……………... 28 Towards a Census of Tropical Rainforest Life: Key outputs ……….… 29 Appendix I. Technical references ……………………………………….. 30 Appendix II. Organizations supporting IBISCA and CTRL …………. 31

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EXECUTIVE SUMMARY

The distribution of Life on Earth Currently, about 1.8 million species are described, of which 80% are arthropods (insects, mites and spiders, the ‘mega-biodiversity’), out of perhaps a total of 5-10 million species on Earth. While there has been substantial effort to database known species, it is fair to say that both description of unknown species in museum collections (aided by new molecular techniques such as DNA barcoding) and field prospecting of yet unknown species lag behind this effort. Biodiversity losses due to human activities were more rapid in the past 50 years than at any time in human history and are projected to continue or accelerate. Further, most of the key outstanding questions in our understanding of biodiversity on Earth relate to beta-diversity – that is: not what exists in the way of genomes, species and ecosystems at a particular place and time but how this changes through both space and time. The nature and mechanisms of species turnover from place to place is one of the major challenges in community ecology especially when the biodiversity being targeted are the massively diverse invertebrates. Tropical rainforests are the most diverse of terrestrial ecosystems and it is accordingly not surprising that efforts to map and understand biodiversity have focused on these forests. It is fair to say that before 2003 most of these studies have been restricted either taxonomically or seasonally, due to the inability to bring sufficient specialized expertise to bear on well-defined contrasting sites.

The significance of tropical forests and their canopies

Tropical forests and their canopies sustain countless species of animals and plants, the majority representing an unknown and unexploited resource. It is widely believed that most of the biological activity in forests, particularly in tropical rainforests, is concentrated in the upper canopy and in the soil. The disruption of healthy forests and their canopies is often irreversible and leads to two obvious perils: (i) loss of forested habitats and concomitant loss of biodiversity and genetic resources; and (ii) loss of ecosystem process such as sequestration of carbon from the atmosphere and concomitant loss of ability to buffer the effects of changes in local and global climates. Against this background, the scientific study of tropical rainforests, from the ground to the canopy, has been impeded by serious difficulties in accessing the canopy. Slowly, the scientific community realizes that the functioning of forest ecosystems cannot be well understood without studying in depth forest canopies.

How to study biodiversity in tropical rainforests: a new approach

We have seen large-scale studies in the tropics before but these have seldom been coordinated against an experimental design aimed at portraying the spatial distribution of organisms against key environmental factors. This began to change with the ‘IBISCA’ project in Panama, initiated in 2003. The project ‘Investigating the Biodiversity of Soil and Canopy Arthropods’ (IBISCA) integrates concerns of spatial, temporal and taxonomic replication. The project studies the beta diversity and vertical stratification of arthropods in the San Lorenzo rainforest in Panama, using state-of-the art methods of canopy access and sampling, namely canopy fogging, canopy cranes, single rope techniques and canopy raft and peripherals. The novelty of the IBISCA project in Panama is that it brought together almost the full set of scientists working globally in the field and recruited them all to a single project. This unprecedented scale of collaboration is generating data sets and understanding at a level of completeness and authority that we have not seen before. For the first time, information on rainforest arthropods has been generated for the whole of the forest (different sites and soil to canopy approach). Despite this, funding for IBISCA has been modest (ca USD300K). IBISCA has set a new ‘industry standard’ involving team work (taxonomists, ecologists, students, parataxonomists), international collaboration and complimentary skills, both in the field and laboratory. In short, the IBISCA project is by far the largest and most comprehensive attempt to date to answer one of the key questions concerning our understanding of the patterns of biodiversity within forest ecosystems. This scientific effort should not stop here.

Why bother? – The applied significance of a project such as IBISCA Why should we bother about preserving countless small invertebrates? These “little things that rule the world” are, together with humans in recent times, the movers and shakers of the world, tearing the vegetation down, cutting paths through the forest, and consuming most of the energy. Invertebrates are key regulators of ecosystem processes, such as herbivory, pollination, seed predation, seed dispersal, decomposition and soil formation. The truth is that we need invertebrates but they don’t need us. If invertebrate species were to disappear, most fishes, amphibians, birds, mammals, flowering plants and, ultimately, the human species would crash to extinction. The applied objectives of a scientific project such as IBISCA are ultimately to prevent such scenario by developing protocols and tools to establish baseline data and quantify the impact of human perturbation, such as loss/fragmentation of forests and global climatic change, on biodiversity. This approach is fully congruent with the scientific pursuit of mapping patterns of biodiversity within tropical rainforests. Further, given the absence of any consensus on the appropriate choice of ‘indicator’ or ‘umbrella’ species, the impacts of human perturbation on biodiversity may be best understood by considering ‘predictor sets’ based on invertebrates, using data generated by projects such as IBISCA.

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A census of tropical rainforest life A proper ‘Census of Tropical Rainforest Life’ (CTRL) would include censuing and studying animals and plants at different heights in most tropical rainforests, a near impossible task. Therefore, we propose to census and study arthropods and vascular plants at different heights within rainforests from key tropical locations, a venture which is within the expertise of the IBISCA group. Such a census would consider about 60-80% of all existing organisms and essentially would aim at understanding the distribution of life on Earth. This census can be modeled after an improved IBISCA protocol. It should be implemented in four incremental steps including a series of pilot studies: (1) Complete all work related to the IBISCA project performed in Panama (completed by mid-2006). (2) Disseminate the important results relevant to the above project and promote the study of mega-biodiversity in the canopies of tropical rainforests (most completed by 2007 but ongoing until 2010). (3) Pave the way for a large international census by completing a series of pilot studies (2006-2010) to develop protocols and tools necessary to initiate CTRL. In particular, some project will focus on the relations between biodiversity and ecosystem function, disturbance and climate change. (4) Eventually, IBISCA would develop an dynamic programme aiming at censuing canopy life at key locations in the tropics, based on lessons learn from (3) (on-going from 2011). Item (1) will include completing the acquisition of ecological variables, taxonomic analyses and databasing, as well as improving the current IBISCA database. Item (2) will be achieved in organizing the current IBISCA group (ca 100 scientists) into a network that will run a single web site and disseminate results efficiently. In particular, a richly illustrated, science-driven, IBISCA book free or sold at a nominal fee could achieve much in promoting the study of canopy mega-biodiversity among experienced as well as future researchers. Item (3) will need to address the following points: maximize current canopy access for biodiversity studies and improve it; maximize the value of the biodiversity information collected in the field and processing of related samples; maximize surveys of arthropods in different tropical habitats; maximize collaboration with existing tropical inventories; assess the relations between biodiversity and ecosystem functions/service; and consider applied aspects of biodiversity research.

Budget and key outputs It is crucial to fund adequate support to process large biodiversity samples (parataxonomist help) and taxonomical expertise (including DNA barcoding), as well as to train PhD and MSc students. The total budget envisioned to achieve activities (1) to (3), above, would be ca US$9.4 million, with US$4.5 million already secured and represented by the salaries of IBISCA experts. The remaining US$4.9 million are sought from institutions and private donors. The outputs of IBISCA and of IBISCA-style pilot studies leading to CTRL are considerable:

• Increased knowledge of the biotic and genetic diversity of the richest habitat on Earth; • Better understanding of key forest processes, such as pollination, herbivory, seed dispersal and decomposition; • Improved conservation policies for the largest share of biodiversity, tropical arthropods; • Dissemination of considerable scientific data: multi-authored paper in high profile journals, a collective book richly

illustrated but free of copyright available in print and downloadable from the IBISCA web site, and a series of more specialized papers;

• Considerable media coverage of IBISCA in the 18 countries from which the IBISCA participants recruit; • A richly illustrated web site which becomes an information source for confirmed canopy researchers, students and the

public, thus promoting media interest and scientific research in tropical forest canopies; • Arthropod collections enriched from material collected in situ in tropical rainforest canopies and ready to be barcoded or

analyzed for genetic content and potentially new molecules; • Numerous new species for science, some which could be named after sponsors; • Capacity building, including exposure to different protocols for confirmed researchers; formation of students in host

countries; and training of parataxonomists which may provide a collateral education in conservation for local communities.

• Applied objectives: developing standard protocols and tools to establish baseline data and quantify the impact of human perturbation, such as loss and fragmentation of forests and global climatic change, on biodiversity.

• Interests for sponsoring: (a) brand name and/or logo on technical devices used to access the canopy, on IBISCA printed books, documents and web site; and (b) positive public image associated with the exploration of the most exciting and rich habitat on Earth.

To our knowledge, there are no alternatives to IBISCA-CTRL if the scientific community is serious about studying tropical arthropods before habitat loss and global climate change extinguish most of this mega-biodiversity.

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RATIONALE Background The distribution of Life on Earth. Without doubt, the most fundamental questions in biology is “How many species are there on Earth” [reference 20 in Appendix I]? A related, equally vital but perhaps methodologically easier investigation is to assess where the greater part of this biodiversity is located. In recent years, there has been considerable debate as whether most of biodiversity occurs in the canopy or in the soil of tropical rainforests [1]. The challenge is to replicate surveys on an appropriate wide range of spatial and temporal scales using a wide variety of sampling protocols for both the canopy and the forest floor. Status of biodiversity on Earth. A recent report [21] makes clear that: (1) loss in biodiversity due to human activities were more rapid in the past 50 years than at any time in human history; (2) the most important drivers of biodiversity loss are habitat change (including loss and fragmentation of forests) and climate change; (3) in the tropics, habitat change contributes much more important to biodiversity loss than climate change do and this situation will continue for a significant time period; and (4) rates of biodiversity loss are projected to continue or accelerate. Currently, about 1.8 million species are described, of which 80% are arthropods (insects, spiders & mites), out of perhaps a total of 5-10 million species on Earth [12, 21]. More biotic inventories are badly needed to refine these estimates, particularly for invertebrates of tropical forest canopies, which represent the largest share of undescribed biodiversity [6, 12, 21]. Biodiversity inventories. Against this background, how has the cataloguing of Life on Earth progressed? While there has been substantial effort to database known species (e.g., Species 2000, Ecoport), it is fair to say that both description of unknown species in museum collections (aided by new molecular techniques such as DNA barcoding [10]) and field prospecting of yet unknown species lag behind this effort. Regarding the latter, a pioneering attempt to launch an “All Taxa Biological Inventory” in Costa Rica [26] failed because of internal dissension, yet survives in a more taxonomically restricted form (ACG, [14]). Other inventories of substantial scope are under way in the tropics (ALAS: [19]; BRC: [22]), the USA (www.discoverlife.org) and Sweden (www.artdata.slu.se/Svenska_artprojektet _Eng.htm). Similar efforts in diverse habitats other than tropical rainforests are proceeding much more rapidly, such as “The Census of Marine Life” (http://www.coml.org/coml.htm). Forest canopies. Further, the scientific study of tropical forests, from the ground to the canopy, has been impeded by serious difficulties in accessing the canopy [18]. The entomologist Terry Erwin labeled tropical rainforest canopies to be ‘The Last Biotic Frontier” on Earth [9]. Progresses have been made in using individual access to the lower part of the canopy (single rope techniques and professional tree climbers), fixed structures such as canopy cranes (www.stri.org/english/research/facilities/terrestrial/cranes/index.php) or mobile devices such as the canopy raft, canopy sledge or tree bubble (www.radeau-des-cimes.org/; Fig. 1). In particular, the cranes of the Smithsonian Tropical Research Institute (STRI), in Panama, were instrumental in facilitating a series of ecological and ecophysiological studies which eventually led to the formation of the International Canopy Crane Network [2]. Slowly, the scientific community realizes that the functioning of forest ecosystems cannot be well understood without studying in depth forest canopies, as recent articles in the leading scientific journal Science indicate [23]. These pressing concerns led the Global Environment Fund and the United Nations Environmental Programme to fund the implementation of canopy observatories which will be activated during 2007-2010 [23]. This represents a global effort to reach the 2010 goals of the United Nations Convention on Biological Diversity, namely to accelerate the inventory of biodiversity on Earth and to slow down the massive genetic extinctions ensuing from the demise of tropical rainforests and global climate change. More than numbers. Most of the outstanding questions in our understanding of biodiversity on Earth relate to beta-diversity – that is: not what exists in the way of genomes, species and ecosystems at a particular place and time but how this changes through space and time. The nature and mechanisms of species turnover from place to place are one of the major challenges in community ecology. Tropical rainforests are the most diverse terrestrial ecosystems and most efforts to map and understand biodiversity have focused on them [4]. It is fair to say that before 2003 and project IBISCA (see below), most studies were restricted either taxonomically or seasonally, because of the inability to bring sufficient expertise together to study well-defined, contrasting sites. Tropical forest and their canopies sustain countless species of animals and plants, the majority representing an unknown and unexploited resource. This reservoir of genetic diversity ensures that vital ecological processes are performed by a variety of species, rather than a few, thus maintaining the integrity of the forest ecosystem. It is widely believed that most of the biological activity in forests, particularly in tropical rainforests, is concentrated in the upper canopy, in comparison with the understorey. The disruption of healthy forests and their canopies is often irreversible and leads to two obvious perils: (i) loss of forested habitats and concomitant loss of biodiversity and genetic resources; and (ii) loss of ecosystem process such as sequestration of

http://www.artdata.slu.se/Svenska_artprojektet _Eng.htm

http://www.coml.org/coml.htm

http://www.radeau-des-cimes.org/

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carbon from the atmosphere and concomitant loss of ability to buffer the effects of changes in local and global climates. Loss of species diversity itself has also direct and serious implications for the maintenance of the functioning of ecosystems. Horizontal distribution (site-to-site), vertical stratification and seasonality are among the most important sources of variance of tropical rainforest diversity [4]. Quantifying these different aspects of diversity for a diverse array of arthropod taxa will in turn improve our understanding of three ecological issues of applied significance: (1) the relations between biodiversity and ecosystem services (forest processes); (2) the impact of disturbance and climate change on biodiversity and forest processes; and (3) biological monitoring. One pressing goal is to understand the distribution of organisms and their dynamic relations with forest processes [21]. Of the initiatives listed above to study biodiversity, only the ACG and BRC projects embrace this approach. However, they are of limited taxonomic focus. Invertebrate contribution to forest processes, such as herbivory, pollination, seed dispersal, decomposition, soil formation, can only be quantified after consideration of a wide array of taxa, including different feeding guilds. Similarly, the magnitude and extent of disturbance can only be assessed by analyzing multitrophic scenarios.

Fig. 1. Methods of canopy access. From top left, clockwise: canopy crane (view of the gondola), canopy bubble, canopy fogging, Ikos (tree platform), single rope techniques and canopy raft. All of these methods were used in the IBISCA project in Panama (see below).

The IBISCA project in Panama We have seen large-scale studies in the tropics before [11, 28; review in 4] but these have seldom been coordinated against an experimental design aimed at portraying the spatial distribution of organisms against key environmental factors. This began to change with the ‘IBISCA’ project in Panama, initiated in 2003. The project ‘Investigating the Biodiversity of Soil and Canopy Arthropods’ (IBISCA) integrates concerns of spatial, temporal and taxonomic replication (public site at http://www.naturalsciences.be/cb/ants/projects/ibisca_main.htm; [7, 24]). It is an initiative of STRI and of Pro-Natural International with technical support of Océan vert. The project studies the beta diversity and vertical stratification of arthropods in the San Lorenzo rainforest in Panama, using state-of-the art methods of canopy access and sampling, namely canopy fogging, canopy cranes, single rope techniques and canopy raft and peripherals. The full range of canopy access techniques is essential to be able to replicate studies of these sites at a wide range of sites (Fig. 1). The Patron of IBISCA is Prof Edward O. Wilson (Harvard University), who is credited with coining the word ‘biodiversity’.


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IBISCA field studies concentrated on spatial replication in 2003 (8 sites surveyed) and temporal replication (4 replicates) in 2004. With funding from Solvin/Solvay, STRI, UNEP, European Science Foundation (ESF) and Global Canopy Programme, IBISCA has studied the arthropod fauna of 8 sites of 400m2 within the San Lorenzo Protected Area, during four study periods: September/October 2003, February/March 2004, May 2004 and October 2004. The canopy was accessed and sampled using a variety of innovative methods. At all sites, the entomology team relied on 14 sampling programs to survey the arthropod fauna of the canopy and of the soil. These sampling programs generated considerable numbers of arthropods, which were sorted with the help of Panamanian students and parataxonomists. Currently, this material is being processed, identified, curated and databased. The entomologists participating in the field work are seconded by many other colleagues for the on-going taxonomic study of the material collected. Each participant concentrates on studying the taxonomy, vertical distribution and beta diversity of particular focal taxa. The vital data will focus on ca. 60 focal taxa, representing a wide array of taxonomic diversity and life histories. Databasing of this information is eased by a modular database developed specifically for the project, available to participants through a password-protected web site. As of September 2005, the IBISCA database which summarizes horizontal (beta diversity), vertical (vertical stratification) and seasonal distribution of arthropods in the San Lorenzo forest included 87 contributors (ecologists and taxonomists), 14 sampling programmes, 7,233 samples, 315 plant species, 428,747 specimens and 1,960 arthropod species. We estimate that the final product should shed light on the spatial and seasonal distribution of about half a million of specimens and several thousands of species, distributed among ca 60 focal groups of different phylogeny and ecology. This dataset has no equivalent. The key questions targeted are: (1) What is the relative contribution of vertical stratification, seasonality and degree of beta-diversity to the distribution of arthropod biodiversity in a closed-canopy tropical rainforest? (2) How do life history traits of species, such as host specificity or feeding guild, influence the spatial and temporal partitioning of arthropod biodiversity in this forest? (3) What can the results tell us about biodiversity estimates, assessment and forest processes?

Fig. 2. Operating standards in the IBISCA project. From top left, clockwise: concentration of biodiversity experts, here surveying insect pollinators in the canopy, at a single location to answer key questions; use of a wide range of collecting techniques, here a flight interception trap, and sufficient taxonomic, spatial and seasonal replicates; pre-sorting of the material into focal taxa involving both students, parataxonomists, ecologists and taxonomists; species identification; databasing using individual modules; and early dissemination of results, such as the vertical stratification of different taxa as collected with sticky traps.

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Preliminary results suggest that (1) despite the short distance between study sites (< 2km), arthropod abundance differ greatly between sites and these differences may be related to forest productivity; (2) faunal overlap is low along the vertical profile of the forest; (3) the higher taxonomic composition of the fauna varies much more between vertical heights than horizontally between sites; (4) arthropod abundance follows a bimodal distribution, with greatest abundance near the ground and in the upper canopy; and (5) patterns of stratification differ among taxonomic and ecological groups, with some being ground- or canopy-dominant. This massive effort caught the attention of the media and of the wider scientific community. So far, the IBISCA project has been presented in 6 scientific articles, 13 magazine articles (written in 8 different languages) and 3 videos but we expect that most of IBISCA results should be disseminated during 2006-2008. A workshop co-funded by ESF and UNEP (see below) and two other symposia at international meetings have provided opportunities to IBISCA participants to review progresses and impediments. The IBISCA workshop in Brussels and the future of rainforest studies An IBISCA workshop was organized at the Royal Belgian Institute of Natural Sciences, in July 2005 and attended by 37 scientists from 16 countries (www.sciencesnaturelles.be/cb/ants/meetings/esf_exploratory_workshop.htm). Its aims were (a) to summarize what has been learned from the IBISCA project overall; (b) to plan meta-data analyses and the dissemination of this novel and important information (see one example in Fig. 3); and (c) to use this material as a springboard to initiate new collaborative programs of research about the distribution of mega-biodiversity in tropical rainforests and to plan a ‘Census of Tropical Rainforest Life’ (CTRL). Given the collective expertise which was present at the Brussels workshop, it is not an exaggeration to state that the near future of canopy biodiversity studies was decided on at this meeting. All workshop participants thought that timing was perfect timing to start developing CTRL, that is, an ambitious program and network dedicated to the proper survey of biodiversity in tropical rainforests. IBISCA has demonstrated that motivated biodiversity experts, even with modest funding, are able to ‘think big’ when studying tropical forest diversity. Expanding projects like IBISCA to regional and global levels to launch CTRL will require a significant lobbying effort for increased funding, but it can be done. Moreover, such new approach to collaborative work in tropical biology and conservation represents an important path to involve local scientists in highly qualified international research. This, in turn, represents a crucial step towards raising the standard of student training and capacity building. The integration of the academia of tropical countries in biological conservation is vital for the study of tropical ecosystems.

Fig. 3. Mean number of arthropods collected per sticky trap at different heights in the San Lorenzo forest during IBISCA. These results are based on one of the 14 sampling programmes used in IBISCA.

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IBISCA started as a relatively modest project in Panama, with limited participants, funding (core project ca $300K) and opportunities to extend it as to cover a substantial part of arthropod diversity and address the effects of seasonality. Due to the great team spirit and the enthusiasm of participants, IBISCA later grew into a full-scale international project. In the field, IBISCA has demonstrated the value of an integrated project to survey tropical arthropods and has provided answers as to how to tackle the study of this mega-diversity efficiently. In short, the IBISCA project is by far the largest and most comprehensive attempt to date to answer one of the key questions concerning our understanding of the patterns of biodiversity within forest ecosystems. This scientific effort should not stop here. Why bother? – The applied significance of a project such as IBISCA Why should we bother about preserving countless small invertebrates? These “little things that rule the world” are, together with humans in recent times, the movers and shakers of the world, tearing the vegetation down, cutting paths through the forest, and consuming most of the energy [29]. Invertebrates are key regulators of ecosystem processes, such as herbivory, pollination, seed predation, seed dispersal, decomposition and soil formation. The truth is that we need invertebrates but they don’t need us. If invertebrate species were to disappear, most fishes, amphibians, birds, mammals, flowering plants and, ultimately, the human species would crash to extinction. The applied objectives of a scientific project such as IBISCA are ultimately to prevent such scenario by developing protocols and tools to establish baseline data and quantify the impact of human perturbation, such as loss/fragmentation of forests and global climatic change, on biodiversity. This approach is fully congruent with the scientific pursuit of mapping patterns of biodiversity within tropical rainforests. Further, given the absence of any consensus on the appropriate choice of ‘indicator’ or ‘umbrella’ species, especially in the tropics [16], the effects of human perturbation on biodiversity may be best understood by considering early ‘predictor sets’ based on invertebrates, using data generated by projects such as IBISCA [15]. Because of their short life cycle and their high sensitivity to environmental changes arthropods can be used as early indicators of ecosystem changes before that effects become apparent on long-lived organisms such as forest trees or mammals. Early predictor sets comprise taxa representative of different functional groups (‘guilds’) as an alternative to the use of taxa selected on the basis of taxonomic tractability or familiarity. Such early ‘predictor sets’ are properly selected only following statistical analysis of a larger, more or less complete, data set including all taxa and the catches from several complementary sampling methods, such as in the project IBISCA [3]. There are also political issues to consider. The United Nations Convention on Biological Diversity (CBD; www.biodiv.org) has committed to achieve by 2010 a significant reduction of the current rate of biodiversity loss [25]. In particular, the Convention specifically calls for a greater emphasis on the forest canopy and the investigation of its interface with the atmosphere. This has created a mandate for governments to support canopy research and is a highly significant development for canopy science and conservation. The text of interest reads as follows [27]: Programme Element 3, Goal 3, Objective 1, Activity b. “Develop and support research to understand critical thresholds of forest biological diversity loss and change, paying particular attention to endemic and threatened species and habitats including forest canopies”. IBISCA represents the largest network of biodiversity experts studying rainforest canopies. Further, the Secretariat of the CBD has identified several major challenges to implementing the convention [27]. Among those, the following items represent particular areas of expertise of IBISCA: (a) increasing the capacity to document and understand biodiversity, its value, and threats to it; (b) building adequate expertise and experience in biodiversity planning; and (c) improving education and public awareness about the value of biodiversity. In particular, taxonomic studies that increase knowledge about the species composition of forests, such as in the proposed programme, are a direct contribution to the activity on forest biodiversity which is part of the programme of work of the Global Taxonomy Initiative (GTI; see Decision VI/8, annex, planned activity #8 of the Conference of the Parties of the Convention of Biological Diversity, available at http://www.biodiv.org). Further, the databases resulting from the programmes will be complied with formats interoperable with major global taxonomic information systems. As such the proposed programme could also contribute to the coordinated global taxonomy information system envisioned in the programme of work of the GTI (planned activity #7).

http://www.biodiv.org/

http://www.biodiv.org/

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Who is IBISCA, why is it different and who supports it? IBISCA is currently an informal group of about one hundred professional scientists, interested in tropical biodiversity and recruiting from the fields of entomology, botany, ecology, taxonomy, statistics and computer science. These scientists belong to 79 institutions based in 18 countries. They will be organized as a formal network shortly. Their joint motivation is to finish the present IBISCA project in Panama and to start developing an ambitious CTRL, based on the lessons learnt from IBISCA. The assets of IBISCA are the collective expertise of all members, which can be estimated as being over 1,500 scientific articles published on tropical biodiversity, 70 books, 500 years spent researching tropical biodiversity, 120 scientific meetings or symposia organized, over 300 PhD and MSc students supervised, over 1,500 new species discovered for Science and over $13.5 million of funds raised to study tropical biodiversity. In short, this is the best network of tropical biodiversity experts. The salaries of IBISCA experts are secured by their respective institutions and amount to $4.5 million for the period covered by this proposal (i.e., more than half of the total costs, see below). The novelty of IBISCA is that it brought together almost the full set of scientists working globally in the field and recruited them all to a single project. Further, the IBISCA approach aims to motivate scientists to work on integrative projects, and is less an inventory per se. For the first time, information on rainforest arthropods has been generated for a wide range and taxa and the whole of the forest (different sites and soil to canopy approach). IBISCA has set a new ‘industry standard’ involving team work (taxonomists, ecologists, students, parataxonomists), international collaboration and complimentary skills, both in the field and laboratory (Fig. 2), and aiming at understanding the distribution of life on Earth. This unprecedented scale of collaboration is generating datasets and understanding at a level of completeness and rigor that we have not seen before. The IBISCA initiative is supported by many academic institutions (NMNH in Washington, NHM in London, STRI in Panama, etc.), research groups (D. H. Janzen, ALAS, etc.) and international networks (CBD, Ecoport, CBOL, GTI, UNEP, etc.). A full list is given in Appendix II.

Fig. 4. View of the canopy of the San Lorenzo forest, home of the IBISCA project.

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TOWARDS A CENSUS OF TROPICAL RAINFOREST LIFE: SCHEDULE AND IMPLEMENTATION

A proper CTRL would include census and study of all prokaryotes, animals, plants and fungi at different heights in most tropical rainforests, a task barely achievable given current levels of expertise, technology and funding [30]. Therefore, we propose to census and study arthropods and vascular plants at different heights within rainforests from key tropical locations, a scheme within the expertise of the IBISCA team. This census can be modeled after an improved IBISCA protocol. Leading topic. The scientific rationale will focus on quantifying the spatio-temporal distribution of a diverse array of focal taxa and relates this information, along with manipulative experiments performed in pilot studies, to applied issues such as (1) the relations between biodiversity and forest processes; (2) the impact of disturbance and climate change on biodiversity and forest processes; and (3) biodiversity monitoring. Our nil hypotheses are that biodiversity is randomly distributed in tropical rainforests (horizontally, vertically and seasonally), that it does not significantly influence forest processes and that disturbance does not significantly impact those. Novel approaches. Most studies of anthropogenic disturbance on terrestrial invertebrates in the tropics are restricted to a handful of groups, not particularly speciose (e.g., [13]). This approach may be overly restrictive given the absence of any consensus on the appropriate choice of ‘indicator’ species and sampling protocols [16,17]. Members of our team have advocated the use of ‘predictor sets’ which comprise taxa representative of different functional groups as an alternative to the use of taxa selected on the basis of taxonomic tractability or familiarity [8, 15]. Such ‘predictor sets’ are properly selected only following statistical analysis of a larger, more or less complete, dataset [3]. A novel strategy that relies on the training and input of local parataxonomists may allow a cost-effective selection of predictor sets (e.g., [5]; www.entu.cas.cz/png/parataxoweb.htm). Further, our team is set to collaborate with the Consortium for the Bar Code of Life, which promotes DNA barcoding techniques [10]. This approach may speed up species identification and will be instrumental in matching life history stages, sexes or damaged specimens. Implementation. Our census should be implemented in four incremental steps, including a series of pilot studies: (1) The ‘IBISCA’ brand needs to be retained as it is valuable and should be used in further projects. The IBISCA research group needs to get organized into a network. The new network needs to be provided with substantial funding as to achieve tasks (2)-(4). (2) The IBISCA network needs to quickly complete the IBISCA-Panama project, to disseminate its results and to promote the study of mega-biodiversity in the canopies of tropical rainforests (completed by 2007, ongoing until 2010). (3) The IBISCA network needs to persist with the IBISCA-approach beyond the Panama project and to initiate new biodiversity projects in Panama and elsewhere (2006-2010). These projects represent pilot studies necessary to initiate CTRL. They will lead to the development of new protocols and tools along the following logical progression and are listed below. They need to be discussed and organized during a series of scientific meetings and conferences. (4) Ultimately, IBISCA needs developing an dynamic programme aimed at censusing rainforest life at key locations in the tropics. This census will focus primarily on arthropods and vascular plants, but with, as far as possible, a view to expanding to other groups and collaborative functional studies. Task (3) includes the following projects:

http://www.entu.cas.cz/png/parataxoweb.htm

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• Project 3.1: Organization of pilot studies: Discuss and organize the required pilot studies during a series of scientific meetings and conferences (2006-2010).

• Project 3.2: Maximize current canopy access facilities for biodiversity research: Expand canopy access within and outside canopy crane perimeters (2006).

• Project 3.3: Improve canopy access for biodiversity research:

Development of mobile canopy access methods and means to use them for biodiversity research (2006-2007).

• Project 3.4: Maximize the value of the biodiversity information collected in the field: Refine inventory methods and protocols (2006-2010).

• Project 3.5: Maximize the processing of biodiversity samples:

Development of an international parataxonomist facility (2006-2010).

• Project 3.6: Maximize surveys of arthropods in different tropical habitats: Insertion of arthropod programmes into large multidisciplinary biological surveys (2006-2010).

• Project 3.7. Maximize collaboration with existing biodiversity inventories:

Collaboration to improve the current inventory of the Area de Conservación Guanacaste (2006-2010).

• Project 3.8: Assess the relations between biodiversity and ecosystem functions/services: Development of protocols for studying ecosystem functions and services in tropical rainforests (2006-2010).

• Project 3.9: Applied biodiversity research:

Assess the impact of disturbance and climate change on biodiversity and ecosystem functions (2006-2010). Note that the scientific community is spending much more on the search for extra-terrestrial life (ETI; it may not even exist within our solar system) than on exploring strategies to stem the loss of biodiversity on Earth (Fig. 5). However, countless unknown species await to be discovered in tropical rainforest canopies on Earth. Hence, each dollar or euro invested in a project like CTRL is likely to represents a higher return than investment into ETI projects and will contribute to alleviate biodiversity loss and alteration of ecosystems on Earth. To our knowledge, there are no other alternatives to the above steps if the scientific community is serious about studying tropical arthropods before habitat loss and global climate change extinguish most of this mega-biodiversity. The total budget to achieve activities (1) to (3), above, would be ca US$9.1 million, with US$4.5 million already secured and represented by the salaries of IBISCA experts. The remaining US$4.6 million are sought from institutions and private donors. The scientific modules and relevant budgets are detailed in the ‘budget explanation’ section, below.

Fig. 5. The rainforest canopy (at left) regulates our global climate and shelter a fantastic amount of new molecules and organisms. However its exploration is currently far less funded than the exploration of space (e.g., planet Mars at right).

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TOWARDS A CENSUS OF TROPICAL RAINFOREST LIFE: BUDGET EXPLANATION

Overview and cumulative budget 2006-2010

Most of the funds sought in this proposal relate to the organization of an international network of experts able to collect and analyze complex biodiversity data (including rounding up the IBISCA-Panama project and disseminating its results); and the implementation of crucial pilot studies (cf. earlier section) necessary to develop the tools and protocols to perform a proper CTRL. Each of these modules is themselves organized as a series of sub-projects, which are detailed below. For most of pilot studies, it is crucial to fund adequate support to process large biodiversity samples (parataxonomist help, see below) and taxonomical expertise, as well as to train PhD and MSc students. Further, IBISCA would like to help to build gradually comparative DNA librairies with barcoding techniques. Current costs include $2.50 needed to sequence a specimens and samples of 10 specimens needed for each species ($25.00 per species). As far as possible, we have included funds in the budgets of relevant sub-projects to sequence the equivalent of 360 species, which will be selected among arthropods of particular importance to forest processes. The total budget envisioned to achieve these various activities would be ca US$9.4 million, with US$4.5 million already secured (salaries of IBISCA experts) and an additional US$4.9 million sought from institutions and private donors. The rationale and budget of each sub-project as listed below is detailed in the following pages.

Module / Sub-project Total budget USD PI Duration

1 Rounding up the IBISCA-Panama project 1.1 IBISCA-Panama: processing of ecological variables 15,000.00 Basset 2006 1.2 IBISCA-Panama: taxonomy 66,000.00 Barrios 2006 1.3 Improvement of the IBISCA database 56,250.00 Leponce 2006

2 Dissemination of results

2.1 IBISCA web site 109,375.00 Leponce 2006-2010 2.2 Dissemination of IBISCA results 65,000.00 Basset 2006-2010 2.3 Running costs for the IBISCA network 341,039.00 Basset 2006-2010

3 Pilot studies

3.1 Organization of pilot studies: scientific meetings 217,250.00 Leponce 2006-2010 3.2 Maximize current canopy access facilities for biodiversity research 75,900.00 Basset/Barrios 2006 3.3 Improve canopy access for biodiversity research 362,900.00 Corbara 2006-2007 3.4 Maximize the value of the biodiversity information collected in the field 517,300.00 Barrios 2006-2010 3.5 Maximize the processing of biodiversity samples 638,500.00 Basset 2006-2010 3.6 Maximize surveys of arthropods in different tropical habitats 455,500.00 Corbara 2006-2010 3.7 Maximize collaboration with existing biodiversity inventories 384,500.00 Basset/Janzen 2006-2010 3.8 Assess the relations between biodiversity and ecosystem functions/services 789,500.00 Kitching 2006-2010 3.9 Applied aspects of biodiversity research 824,500.00 Leponce 2006-2010

Grand total 2006-2010 4,918,514.00 Budget 2006 1,263,092.80 Budget 2007 1,049,942.80 Budget 2008 868,492.80 Budget 2009 868,492.80 Budget 2010 868,492.80

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Module 1: Rounding up the IBISCA-Panama project

Sub-project 1.1: IBISCA-Panama: processing of ecological variables Aims. To finish processing information currently missing for the IBISCA-Panama project and help a better interpretation of the data related to this project. Rationale. This sub-project has been identified as crucial to complete the IBISCA-Panama project by all participants of the IBISCA Brussels workshop. The missing ecological variables will greatly help in the final interpretation of IBISCA data. It will include the following items: (1.1.1) processing about 1,000 digital pictures to estimate canopy openness at all IBISCA study sites; (1.1.2) acquisition of high resolution pictures of the San Lorenzo forest to derive indices of forest productivity for all IBISCA study sites; (1.1.3) completion of canopy pin point studies at four IBISCA sites not surveyed during 2003-2004 and get estimates of leaf index and herbivory at these sites; and (1.1.4) estimation of various site variables during a brief field study. Output. A new set of variables available in the IBISCA database.

Item Budget USD PI

1.1.1 Canopy openness: hemispherical pictures 3,000.00 Basset/Roslin/Didham 1.1.2 Acquisition of remote sensing data 3,000.00 Ribeiro 1.1.3 Canopy pin point studies for 4 sites not surveyed during 2003-2004 8,000.00 Ribeiro 1.1.4 Assess disturbance, soil typology and slope of study sites 1,000.00 Basset

Grand total 15,000.00

Fig. 6. One of the thousands of digital pictures to process to get accurate estimates of canopy openness at the different IBISCA study sites in Panama. This picture was shot below a sticky trap (in yellow), collecting small insects in the understorey.

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Sub-project 1.2: IBISCA-Panama: taxonomy Aims. To round-up taxonomical analyses of the material collected during the IBISCA-Panama project. Rationale. The items listed below are crucial to complete specimen processing, identification and databasing, and, over the longer-term, the preparation of a high-profile collective article in a leading journal. This will allow us to: (a) speed up the processing and morphotyping of specimens (Oribatida, Hymenoptera Parasitica; ants, etc.); (b) speed up databasing of specimens (e.g. Ødegaard’s beetle focal taxa); © visit insect collections (Braconidae: Texas; Curculionoidea+Auchenorrhyncha: InBio in San José, etc.); (d) expand the range of focal taxa (e.g., Collembola). Outputs. Arthropod collections (specimens, slides, etc.), description of new species, and improvement of the IBISCA database.

Item Budget USD

PI

1.2.1 Processing and identification of Acari Oribatida 6,000.00 Winchester 1.2.2 Visit to Texas A&M University collections and identification of Braconidae 5,000.00 Medianero 1.2.3 Processing of Hymenoptera Parasitica in Malaise trap samples 5,000.00 Springate 1.2.4 Visits to InBio (Costa Rica) and Ottawa collections and identification of Curculionoidea 3,000.00 Barrios 1.2.5 Identification of Cucujoidea 3,000.00 Cizek 1.2.6 Revision of Agrilus in Panama 3,000.00 Curletti 1.2.7 Identification and databasing of Formicidae 3,000.00 Delabie 1.2.8 Identification and processing of various groups of Diptera 3,000.00 Didham & Fagan 1.2.9 Identification and databasing of various groups of Coleoptera 3,000.00 Odegaard 1.2.10 Identification of various Coleoptera, Araneae and Blattodea 3,000.00 Schmidl 1.2.11 Processing and identification of Nitidulidae 3,000.00 Tishechkin 1.2.12 Processing and identification of Staphylinidae 3,000.00 Winchester 1.2.13 Processing and identification of Ptilodactylidae and Chalcididae 2,000.00 Aberlenc 1.2.14 Identification of Orthoptera 2,000.00 Bail 1.2.15 Visit to InBio (Costa Rica) and identification of Auchenorrhyncha 2,000.00 Basset 1.2.16 Expand the range of focal taxa: Collembola 2,000.00 Castano-Meneses 1.2.17 Identification of Ispopoda and Myriapoda 2,000.00 Cizek 1.2.18 Processing and identification of Chrysomelidae 2,000.00 Floren 1.2.19 Identification of Isoptera 2,000.00 Roisin


3 Wasmannia sp.1

Solenopsis sp.1

1 Brachymyrmexphysogaster

2

4. Paratechina sp.2 5. Octostrumarugifera

6. Pheidole sp.1

7. Crematogastersp.2

8. Hypoponerasp.4

9. Hypoponerasp. prox. trigona

10. Pyramicadenticulata

11. Pheidoleradoszkowskiireflexans

Fig. 7. Some of the ant species collected in the San Lorenzo forest, out of a total of ca 150 species.

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Sub-project 1.3: Improvement of the IBISCA database Aims. To improve and clean the IBISCA database, so that IBISCA members can use it efficiently to report on the main results of the IBISCA-Panama project. Rationale. One of the working group issued from the IBISCA Brussels workshop will implement a common set of statistical analyses (as statistical modules in the IBISCA database) that will allow the IBISCA group to report collectively their results in a leading scientific journal around mid-2006. To this end, this working group should meet in Brussels, where the majority of group participants reside. In addition, a certain number of improvements will be needed for the IBISCA database, namely: (a) determine which plant specimens are located in the understorey/mid-canopy/upper canopy on the basis of regression equations between DBH and height, and calibration in the field; (b) add meteorological and tree phenology data to the database; (c) propose other ways of expressing arthropod dispersal (e.g. geography, distance) and add such a field in the database; (d) add the fields ‘habitat specificity’ and ‘host specificity’ for each species; (e) clean-up of the database. Output. A greatly improved database, which will be useful for pilot studies.

Item Budget USD PI

1.3.1 Database improvement, including programmer 56,250.00 Leponce Grand total 56,250.00

Fig. 8. Screenshot of the opening menu of the IBISCA database.

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Module 2: Dissemination of results

Sub-project 2.1: IBISCA web site Aims. To develop a user-friendly IBISCA web site centralizing all information available on IBISCA projects. Rationale. Ideally, a single IBISCA web site should be implemented (currently two sites are active, one public, one password-protected). This site should include items related to the IBISCA network; the IBISCA database, media reports and scientific articles, etc. Access to different web pages should be either public or password-protected, depending on the sensitivity of the information. As far as possible, the web site should also include digitized pictures of all IBISCA morphospecies. It should display a number of important links, especially taxonomic links and funding sources. A person should be hired to develop the interface, manage the web site and the associated correspondence. Output. An improved web site.

Item Budget USD PI

2.1.1 Implementation of a unique web site, both password-protected and public 10,000.00 Leponce 2.1.2 Measurement of body size and digitized pictures for all morphospecies 50,000.00 Leponce 2.1.3 Running costs and improvements 10,000.00 Leponce 2.1.4 Salary for web site manager (2006-2010) 39,375.00 Leponce


Fig. 9. Opening screen of the IBISCA public web site.

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Sub-project 2.2: Dissemination of IBISCA results Aims. To disseminate the results of the IBISCA-Panama project and promote the study of mega-biodiversity in tropical rainforests. Rationale. Budget for this sub-projected is mainly related to the publication of a collective book free of copyright on the preliminary results from the IBISCA-Panama project. IBISCA has so far raised ca $13,000 towards this end but needs to secure more funds. The project is innovative as it provides in one package:

• First rate science: preliminary results of IBISCA and reports originating from the present workshop. Perhaps 40 multi-authored chapters of 10 pages each, plus references and index.

• Attractive iconography: many colour pictures, perhaps organized in plates (2-3 plates per chapter if possible). • Searchable companion CD. • Public access to downloadable pdf files and additional information (pictures, updated species lists), hosted on the IBISCA

web site. • Free dissemination to targeted readers and groups (scientists and institutions in developing countries: university librairies,

IBISCA participants): perhaps 500 copies. • Nominal fee for other readers: perhaps 500 copies at $5-10 each, to cover mailing expenses associated with the above

item. The final product is aimed as a cross between a standard edited scientific book and a richly illustrated ‘coffee table’ book. This represents one of the best ways to promote the study of canopy tropical biology. In addition, other funds are needed to lower page charges (if applicable) in scientific journals for the publication of the main IBISCA results. Outputs. One collective article in a leading science journal; a collection of refereed and multi-authored articles; and a richly illustrative collective book whose copyright is retained by IBISCA.

Item Budget USD PI Remark

2.2.1 IBISCA collective book (richly illustrated) 30,000.00 Basset $13,000 already available 2.2.2 Book handling and mailing expenses 10,000.00 Schmidl 2.2.3 Creation of html files and CDs related to the book 5,000.00 Leponce 2.2.4 Page fees for main IBISCA articles 20,000.00 Basset


Sub-project 2.3: Running costs for the IBISCA network Aims. To formally organize the current IBISCA research group into an efficient network. Rationale. Results of discussions among IBISCA participants established the need for the IBISCA research group (about one hundred scientists) to get organized into a formal network. This is likely to include secretarial costs, including the implementation of the new network. Half-time salaries are needed for an overall manager and an assistant. Outputs. A formal network, with web site and secretariat.

Item Budget USD PI

2.3.1 Steering committee meetings (n = 5) 37,500.00 Leponce 2.3.2 Manager: 5 years half-time 187,500.00 Basset 2.3.3 Assistant to the manager: 5 years half-time 116,039.00 Missa


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Module 3: Pilot studies

Sub-project 3.1: Organization of pilot studies: scientific meetings

Aims. To plan efficiently the dissemination of IBISCA-Panama data and pilot studies through a series of workshops. Rationale. Planning the organization and protocols of the different pilot studies is necessary and will involve IBISCA participants and representatives of international biodiversity networks/research groups. A series of 9 workshops (3 days, 20-25 scientists) and one final conference (3 days, 50 scientists) will allow a better preparation of the different pilot studies. Outputs. A series of concerted plans for the proposed pilot studies and final guidelines as to how to complete a CTRL. Geographic location: Various, not decided yet.

Item Budget USD PI

3.1.1 Workshop 1: Statistical tools and analyses of biological surveys 17,500.00 Leponce 3.1.2 Workshop 2: IBISCA-Panama: results, dissemination and lessons 17,500.00 Leponce 3.1.3 Workshop 3: Scientific/public outreach, property right issues and funding 17,500.00 Leponce 3.1.4 Workshop 4: Focal arthropod taxa, forest processes, taxonomy, barcoding 17,500.00 Leponce 3.1.5 Workshop 5: Development of an international parataxonomist facility 17,500.00 Leponce 3.1.6 Workshop 6: Improving canopy access for biodiversity research 17,500.00 Leponce 3.1.7 Workshop 7: Refining survey protocols at different tropical locations 17,500.00 Leponce 3.1.8 Workshop 8: Relations between biodiversity and forest processes 17,500.00 Leponce 3.1.9 Workshop 9: Disturbance/climate change and biodiversity/forest processes 17,500.00 Leponce 3.1.10 Final conference summarizing progresses: Towards a CTRL 40,000.00 Leponce Contigency fund (10%) 19,750.00


Fig. 10. One of the working groups during the IBISCA-Panama workshop held in Brussels in July 2005.

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Sub-project 3.2: Maximize current canopy access facilities for biodiversity research Aims. To maximize current canopy access facilities for biodiversity research. In practice this includes expanding canopy access within and outside canopy crane perimeters. Rationale. It is imperative that studies of forest processes account for the biological activity occurring in the upper canopy. Canopy facilities based on canopy cranes are gaining popularity world-wide (10 such facilities currently and 5 more planned by the GEF-UNEP project, see Background). However, access to the lower parts of the canopy is also critical to study vertical gradients in abiotic and biotic factors from the ground level to the upper canopy, and to scale up measurements to the whole-forest level. Access to lower parts of the canopy is difficult with a canopy crane which, furthermore, provides limited spatial replication. A solution is to develop safe and non-intrusive protocols for conducting scientific surveys and experimentation within the upper canopy with a crane, and rope-based canopy access to replicate these observations in the lower parts of the canopy inside as well as outside the crane perimeter. These protocols could be developed through a pilot study at the San Lorenzo canopy crane in Panama and may be later replicated within the existing network of canopy cranes, particularly at tropical locations. The end product is envisioned as improving canopy access at existing canopy facilities, so that IBISCA experts can apply safely their protocols there. This sub-project will include the safe installation of additional canopy access points by two professional tree climbers. Outputs. A model protocol for improving canopy access at existing canopy crane facilities, and one web of access points at the San Lorenzo canopy crane. Geographic location: Panama

Item Budget USD PI

3.2.1 Single rope technique equipment 10,000.00 Global Canopy Programme 3.2.2 Surveying, choosing and setting access points Global Canopy Programme Climber international flights (n = 2) 2,500.00 Climber food and �ccommodation at STRI 1,500.00 Local travel: rental of 4x4+ gas 1,000.00 Salary for climbers 20,000.00 Crane and STRI fees 1,500.00 Insurance 2,000.00 3.2.3 Material for access points (incl. platforms) 15,000.00 Global Canopy Programme 3.2.4 Maintenance of access points 15,000.00 Global Canopy Programme 3.2.5 Report 500.00 Barrios/Basset Contigency fund (10%) 6,900.00


Fig. 11. An IBISCA participant measuring herbivory with single rope techniques at different heights within the San Lorenzo forest

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Sub-project 3.3: Improve canopy access for biodiversity research Aims. To develop mobile canopy access methods and means to use them for biodiversity research Rationale. Access to forest canopies is difficult, impeding efforts to research and to survey a rich biodiversity. Flying over the canopy may be on the best strategies to uncover the secrets of tropical rainforests. In particular, a mobile device may increase replication over spatial scales, as opposed to fixed facilities, such as canopy cranes. To explore the canopy, Pro-Natura and partner Océan Vert have built a toolset based on the Canopy Raft. One of these tools – the Canopy Sledge – was particularly practical for researchers because of its ease-of-use and its mobility. It has inspired the Canopy-Glider, a smaller and more high-performance machine. The Canopy-Glider provides a faster, more reliable method than those currently available for visually locating the positions of the trees or canopy-dependent organisms, recording and mapping them, etc. The Canopy-Glider is a powerful and efficient sampling tool that can also be used to sample and assess genetic variability within or between populations. Low altitude flights can facilitate a wide variety of activities: • Continuous flights can be used for: - aerial photography for canopy tree identification, detailed mapping, etc.; - census of trees, or of other organisms such as large vertebrates (birds, primates, etc.). • Step-by step flights with short stops in the canopy can be used for: - collection of flowers, fruits and leaves of trees and lianas, epiphytes; - periodic sampling and census of canopy dwelling organisms using traps, pre-established stations, etc. • Longer-term stops in the canopy can be used for: - set up of recording stations for biophysical and biochemical parameters; - placement of traps for pollen, air-borne organisms, insects, vertebrates, etc. The Canopy-Glider is being designed to allow 3 persons to move in contact with the canopy for periods of 2 to 3 hours without interruption (usually in the morning or at night when wind velocity is lowest). It will be simple to stop whenever necessary to make observations, gather samples, conduct analysis, photograph or film the canopy, etc. This IBISCA sub-project will perform a mission of the Canopy Glider in French Guiana, led by Bruno Corbara (IBISCA). This contribution would help ca 10 IBISCA experts to implement specific protocols based on the mobility of the Canopy Glider. Outputs. A model mission with the Canopy Glider. Geographic location: French Guiana, possibly Area de Conservación Guanacaste, Costa Rica.

Item Budget USD PI

3.3.1 Model mission in French Guiana with Canopy Glider Corbara Preparation 36,000.00 Daily expenses: $4,580 per day (30 days) 137,400.00 Insurance and freight 72,000.00 3.3.2 Parataxonomist help 25,000.00 Novotny 3.3.3 Taxonomic studies 50,000.00 Barrios 3.3.4 Barcoding of selected species 9,000.00 Barrios 3.3.5 Report 500.00 Corbara Contigency fund (10%) 33,000.00


Fig. 12. The Canopy Glider.

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Sub-project 3.4: Maximize the value of the biodiversity information

collected in the field Aims. To maximize the value of the biodiversity information collected in the field by refining inventory methods and protocols. Rationale. This sub-project will allow refining the survey protocols of the most efficient sampling methods used during the previous IBISCA-Panama project. In particular, we will explore how to optimize sampling towards key focal taxa for molecular analyses (barcoding technique), so that arthropod phylogenies can be compared with geological history. Field work will be based at several locations in Panama, at the request of Authoridad Nacional del Ambiante – Panama and GEF (Héctor Barrios, IBISCA, PI). The isthmus of Panama is of particular interest to evolutionary biologists, as its past geological history may explain in part the contemporary distribution of plants and animals. The sub- project will basically follow-up the general aims of the IBISCA-Panama project (study of the horizontal, vertical and seasonal distribution of arthropods in forests), but will expand them as to cover different types of forests: lowland dry tropical semi-deciduous forest, lowland tropical wet evergreen forest, and montane tropical evergreen forest. A pilot study will be organised at one of the study sites in 2006, then full-scale field work will be replicated in 2007 and 2008 (20 biodiversity experts). Outputs. Development of sound methodology and tools to survey efficiently biodiversity. Geographic location: Panama, including Parque La Amistad, one of the UNESCO-MAB site.

Item Budget USD PI

3.4.1 Three international trips for experts (n=20) 72,000.00 Barrios 3.4.2 Food and accommodation for experts 73,800.00 Barrios 3.4.3 Local travel and gasoline 10,000.00 Barrios 3.4.4 Entomological equipment 59,000.00 Barrios 3.4.5 Field equipment 20,000.00 Barrios 3.4.6 Salaries local assistants during field work(n=10) 12,000.00 Barrios 3.4.7 Facility fees, incl. Canopy cranes 40,000.00 Barrios 3.4.8 Collecting/export permits and mailing expenses 10,000.00 Barrios 3.4.9 Stationary and varia 2,000.00 Barrios 3.4.10 Parataxonomist help (n = 4) for 3 years 57,600.00 Basset 3.4.11 Taxonomic studies 50,000.00 Barrios 3.4.12 Barcoding of selected species 9,000.00 Barrios 3.4.13 Student bursaries (n = 4) for 2 years 54,400.00 Barrios 3.4.14 Report 500.00 Barrios Contigency fund (10%) 47,000.00


Fig.13. The three type of forest that will be investigated in Panama during the GEF-ANAM project, from left to right:

lowland dry semi-deciduous, lowland wet evergreen and montane evergreen forest.

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Sub-project 3.5: Maximize the processing of biodiversity samples Aims. To maximize the processing of biodiversity samples by developing an international parataxonomist facility. Rationale. The lack of primary information on the ecology of most terrestrial arthropods in the tropics appears to be the principal obstacle to the rapid development of practical recommendations for the conservation of rainforest ecosystems. It is crucial that we rehabilitate systematics as a fashionable discipline and increase its global capacity. Parallel efforts should also aim to improve the flow of primary information on tropical biodiversity. In brief, at least a hundred-fold increase in the efficiency of data collection is required if tropical ecology is to proceed beyond its current embryonic stage. This goal is not unrealistic [5]. Data collecting by parataxonomists represents one of the most efficient approaches to the study of tropical ecosystems available to date. As the name implies and was originally defined, parataxonomists stand ‘at the side’ of taxonomists and the taxonomic process. Through iteration, their mostly rural career is to conduct, magnify and improve the processing of specimens and collateral information collection in the field. The expertise of parataxonomists lies in collecting specimens, preparing them, preliminary sorting into morphospecies, and databasing the associated information. Their work results in quality material that can be deposited in national collections and used for taxonomic studies. Although their role is more demanding than that of local informants (e.g. ‘tree-spotters’), they are not an alternative to professional taxonomists in the field or laboratory, but rather enhance their activities and capacities. Further, the indirect, positive effects of local involvement in research should not be underestimated. Involvement of local communities in ecological research affirms the value of biological conservation. Collateral education of local people by fellow parataxonomists may also be significant. It is crucial to consider parataxonomist help to facilitate future IBISCA-type projects in developing an international ‘parataxonomy facility’. We propose to train efficiently some parataxonomists of The New Guinea Binatang Research Center (www.entu.cas.cz/png/parataxoweb.htm) and Area de Conservación Guanacaste (http://janzen.sas.upenn.edu/) to sort and process focal groups of interest to IBISCA. This will include 5 training courses and salaries for 10 parataxonomists during 3 years. Outputs. An efficient team of 10 parataxonomists; arthropod collections; collateral education of local people. Geographic location: Papua New Guinea, Costa Rica and international.

Item Budget USD PI

3.5.1 Parataxonomist training course – Lepidoptera 20,000.00 Kitching 3.5.2 Parataxonomist training course – Coleoptera 20,000.00 Odegaard/Schmidl/Aberlenc 3.5.3 Parataxonomist training course – Hymenoptera 20,000.00 Springate/Delabie 3.5.4 Parataxonomist training course – Diptera 20,000.00 Didham/Rapp 3.5.5 Parataxonomist training course – Other insect orders 20,000.00 Basset 3.5.6 Parataxonomist training course – Other arthropods 20,000.00 Basset 3.5.7 Equipment (computers, microscopes, digital cameras) 50,000.00 Novotny/Janzen 3.5.8 Salaries: 10 parataxonomist full-time for 5 years 360,000.00 Novotny/Janzen 3.5.9 Mailing expenses for specimens 50,000.00 Novotny/Janzen 3.5.10 Report 500.00 Novotny/Basset/Janzen

Contigency fund (10%) 58,000.00 Grand total 638,500.00

Fig. 14. Example of activities involving parataxonomists, clock-wise from left to right: collecting plant and insects in the field;

rearing live insect specimens; mounting insect specimens; and databasing.


http://janzen.sas.upenn.edu/

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Sub-project 3.6: Maximize surveys of arthropods in different tropical habitats Aims. To maximize surveys of arthropods in different tropical habitats and micro-habitats; to insert arthropod programmes into large multidisciplinary biological surveys. Rationale. The IBISCA-Panama project has set a new working standard to survey arthropods in tropical rainforests. However, tropical terrestrial arthropods also thrive in other habitats, such as freshwater, grasslands, caves, etc. It is important to evaluate which proportion of biodiversity is supported by habitats other than rainforests. Further, within rainforests, many specific micro-habitats support an often very specialized fauna. Therefore, it is important to develop protocols and tools to survey arthropods in a variety of macro- and micro-habitats. SANTO is a scientific expedition which will survey the flora and fauna of terrestrial and marine habitats of a large mountainous island in the South Pacific: Espiritu Santo (or Santo), in Vanuatu. Surveys will be implemented for all the main island habitats: deep benthos, coral reefs, caves, freshwater, montains, forest canopies, etc. The project will also consider local knowledge and relationships with scientific knowledge. This will include a considerable effort, including both human personnel (professional divers, cavers, tree climbers, ethnobiologists and biologists) and infrastructure (research vessel, tree glider [see next section]). More than 100 experts from all over the world will be performing field work at Santo. Particular emphasis will be given to the implication of local partners in research, to the restitution of the information for sustainable development, to conservation education, as well to the coordination of national and international research programmes in Vanuatu. The scientific programme includes 5 main ‘modules’, organized around survey techniques, and one theme common to all modules, ‘ethnoscience’. In addition to important collections of specimens, the objectives of the 5 modules will depend on similar questions: what is the real magnitude of biodiversity in the most rich habitats? What is the influence of rare species in floristic/faunistic communities? What is the spatial dimension of biodiversity and what is the representativeness of study sites at the regional scale? The lead institutions for this project are based in France and include the Muséum National d’Histoire Naturelle (MNHN), the Institut de Recherche pour le Développement (IRD) and Pro-Natura International (PNI). Coordination will be under the lead of Philippe Bouchet (MNHN), Pierre Cabalion (IRD), Bruno Corbara (IBISCA), Hervé Le Guyader (IRD, CNRS, MNHN) and Olivier Pascal (PNI-IBISCA). The total budget for the SANTO project is estimated to be ca 1mio of euros. Field work on terrestrial components by ca 20 IBISCA experts during 3 years is estimated to cost $300K, including their use of the Canopy Glider (sub-project 3.2). The IBISCA-SANTO project will represent the ideal opportunity for the IBISCA group to implement a model programme based on surveying a range of habitats and to insert this programme into a multidisciplinary survey, including ethnography, the diversity of arthropods and vegetation, invasive species, tropical cave systems, etc. Outputs. Protocols and tools to survey efficiently a range of macro- and micro-habitats. Geographic location: Santo Island, Vanuatu.

Item Budget USD PI

3.6.1 Field work in Vanuatu, terrestrial component 300,000.00 Corbara 3.6.2 Parataxonomist help 25,000.00 Novotny 3.6.3 Taxonomical studies 50,000.00 Barrios 3.6.4 Student bursaries 30,000.00 Corbara 3.6.5 Barcoding of selected species 9,000.00 Barrios 3.6.6 Report 500.00 Corbara Contigency fund (10%) 41,000.00


Fig. 15. Alluvial forest North of Santo, Big Bay.

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Sub-project 3.7: Maximize collaboration with existing biodiversity inventories

Aims. A collaboration to improve the current inventory of the Area de Conservación Guanacaste (2006-2010). Rationale. A few biodiversity inventories are currently underway in the tropics. They all differ with respects with aims, scope, funding, etc. One challenge is for IBISCA to capitalize on these different projects and to collaborate efficiently with these various research groups as to speed up a proper CTRL. One key tropical inventory is D.H. Janzen’s census of caterpillars feeding on plants of the Area de Conservación Guanacaste in Costa Rica (ACG; Reference [14]; http://janzen.sas.upenn.edu/). The inventory started in 1978 and concerns +4,000 species of macrolepidopterans and their Diptera and Hymenoptera parasitoids. Opportunities for collaboration between ACG and IBISCA are multiple, but need to suit both the interests of ACG and IBISCA. We believe that two avenues are worth pursuing: (a) improvements on the actual inventory as to include forest types and microhabitats currently not covered; and (b) expanding the food webs studied as to include microlepidopterans, mining and galling caterpillars, and perhaps other taxa of immediate relevance to the ACG inventory. Another avenue of collaboration may be explored with the inventories currently being developed by the Binatang Research Center in Papua New Guinea (www.entu.cas.cz/png/parataxoweb.htm). Outputs. Refined protocols, better collaboration and increased capacity to undertake a proper CTRL. Geographic location: Area de Conservación Guanacaste, Costa Rica.

Item Budget USD PI

3.7.1 Module I: contribution to caterpillar inventory 60,000.00 Janzen 3.7.2 Salaries of paratxonomists - module I 60,000.00 Janzen 3.7.3 Module II: expanding food webs 60,000.00 Basset 3.7.4 Salaries of parataxonomists - module II 60,000.00 Janzen 3.7.5 Training course for parataxonomists 20,000.00 Novotny/Janzen 3.7.6 Taxonomic studies 50,000.00 Barrios 3.7.7 Barcoding of selected species 9,000.00 Barrios 3.7.8 Student bursaries 30,000.00 Janzen 3.7.9 Report 500.00 Basset


Fig. 16. Caterpillars collected in the upper crown of trees with a cherry-picker vehicle, in the ACG.

http://janzen.sas.upenn.edu/


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Sub-project 3.8: Assess the relations between biodiversity and ecosystem functions/services

Aims. To assess the relations between biodiversity and ecosystem functions/services and to develop protocols for studying ecosystem functions and services in tropical rainforests Rationale. Relationships between biodiversity and forest ecosystem functions, such as herbivory, pollination, seed dispersal and decomposition, may be better understood by contrasting changes in forest function and fauna along environmental gradients, such as altitudinal gradient. The IBISCA-BATH project in Australia which will focus on the study of such a gradient, will therefore provide an ideal opportunity to develop specific protocols to study forest ecosystem functions. The IBISCA-BATH project, modelled on the successful IBISCA Project conducted in Panama, will bring together multinational teams of biodiversity specialists to quantify biodiversity changes over a 1000m altitudinal gradient in the subtropics of eastern Australia. It is proposed that there be two four-week periods of field work (ca. October 2006, April 2007) during which specialists in a set of target taxa of arthropods, plants and fungi will carry out surveys at 24 sites arrayed at 200m altitudinal intervals along the transect in south-east Queensland. The Project will be led by Professor Roger Kitching (Griffith University and IBISCA), assisted by an executive group from the Queensland Museum, Queensland Herbarium, Queensland Department of Premier and Cabinet, Queensland National Parks and Wildlife Service and the National Parks Association of Queensland. The following main hypotheses will be investigated: 1. There are changes in the faunal and floristic assemblages which mirror altitudinal change within a continuous area of rain forest. 2. There are measurable changes in the rates and nature of key ecological structures and processes – decomposition, food-web structure, herbivory and predation – across altitudinal gradients within a forest. 3. The overall, altitudinally oriented set of results will represent a powerful monitor of future climatic (or other ecosystem-level) change which will be detectable through small scale focused surveys based on the overall results we shall obtain. The conceptual framework for the project will be to target a set of focal taxonomic groups spanning a wide range of functional roles within the rainforest ecosystem. All compartments of the forest at any one location, from the soil to the canopy, will be examined. Outputs. Specific protocols and tools to assess the relation between biodiversity and forest functions. Geographic location: Australia, Lamington Park (Queensland).

Item Budget USD PI

3.8.1 Travel (domestic and international) 105,000.00 Kitching 3.8.2 Accomodation & board 180,000.00 Kitching 3.8.3 Equipment & shipping 270,000.00 Kitching 3.8.4 Student bursaries 8,000.00 Kitching 3.8.5 Administrative costs 40,000.00 Kitching 3.8.6 Module herbivory 10,000.00 Kitching 3.8.7 Module pollination/seed dispersion 10,000.00 Kitching 3.8.8 Module decomposition 10,000.00 Kitching 3.8.9 Parataxonomist help 25,000.00 Novotny 3.8.10 Taxonomic studies 50,000.00 Barrios 3.8.11 Barcoding of selected species 9,000.00 Barrios 3.8.12 Report 500.00 Kitching


Fig. 17. View of Lamington Park, near Brisbane, Australia, home of the BATH project.

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Sub-project 3.9: Applied aspects of biodiversity research

Aims. To assess the impact of disturbance and climate change on biodiversity and ecosystem functions. Rationale. The objectives of this sub-project would be met in three steps: (a) exploring the data of IBISCA sub-projects 3.1 to 3.7; (b) organizing two workshops, with many IBISCA experts, to plan field work and monitoring related to disturbance (forest loss and fragmentation) and global climate change, respectively; and (c) perform two field studies, including monitoring periods, related to disturbance and climate change, respectively. The disturbance study will be set in Panama, to take advantage of the data collected during the IBISCA-Panama projects, as well as sub-projects 3.1 and 3.3. The climate change study will be performed in Lamington Park, Australia, to expand on the data collected with sub-project 3.8. Outputs. Specific tools to assess the impact of disturbance and climate change on biodiversity and ecosystem functions. Geographic location: Panama and Australia, with the Area de Conservación Guanacaste, Costa Rica, another possibility.

Item Budget USD PI

3.9.1 Exploring data of previous IBISCA projects 10,000.00 Leponce 3.9.2 Field work-monitoring: disturbance 350,000.00 Basset 3.9.3 Field work-monitoring: climate change 350,000.00 Kitching 3.9.4 Student bursaries 30,000.00 Kitching 3.9.5 Barcoding of selected species 9,000.00 Barrios 3.9.6 Report 500.00 Leponce Contigency fund (10%) 75,000.00


Fig. 18. Results of selective logging in Guyana. From top left, clockwise: undisturbed understorey; large natural forest gap; selective logging; result of intense selective logging.

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TOWARDS A CENSUS OF TROPICAL RAINFOREST LIFE: KEY OUTPUTS

Outputs specifically related to IBISCA-Panama: • Better understanding of the spatial, vertical and temporal distribution of rainforest arthropods; • Dissemination of considerable scientific results based on the IBISCA-Panama data: multi-authored paper in high profile

journals, a collective book richly illustrated but free of copyright available in print and downloadable from the IBISCA web site, and a series of more specialized papers authored by the participants;

• A scientific database summarizing the distribution of arthropods in the San Lorenzo forest with no equivalent;

Outputs related to pilot studies leading to CTRL • Increased knowledge of the biotic and genetic diversity of one of the richest habitat on Earth; • Better understanding of key forest processes, such as pollination, herbivory, seed dispersal and decomposition; • As a result of above items, better conservation policies related to the largest share of biodiversity, tropical arthropods; • An renowned team of biodiversity experts formally organized in a research group, ready to tackle a ‘Census of Tropical

Rainforest Life’; the collective experience of these experts has no equivalent; • Considerable media coverage of IBISCA in the 16+ countries from which the IBISCA participants recruit; • A richly illustrated web site which becomes an information source for confirmed canopy researchers, students and the

public, thus promoting media interest and scientific research in tropical forest canopies; • Arthropod collections enriched from material collected in situ in tropical rainforest canopies and ready to be bar-coded

or analyzed for genetic content and potentially new molecules; • Numerous new species for science, some which could be named after sponsors; • Development of key solutions to achieve quickly a global inventory of Life on Earth: improved canopy access at

existing canopy facilities; local parataxonomist training, mobile access to forest canopies, improved survey protocols well integrated which allow studying ecosystem functions and evolutionary scenarios.

• Capacity building, including exposure to different protocols for confirmed researchers; formation of students in local and host countries; and training of parataxonomists which may provide a collateral education in conservation for local communities.

• Applied objectives: developing standard protocols and tools to establish baseline data and quantify the impact of human perturbation, such as loss and fragmentation of forests and global climatic change, on biodiversity.

• Interest for sponsoring: (a) brand name and/or logo on technical devices used to access the canopy, on IBISCA printed books, documents and web site; and (b) positive public image associated with the exploration of the most exciting and rich habitat on Earth.

Fig. 19. Key activities during the project IBISCA-Panama

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APPENDIX I. TECHNICAL REFERENCES 1. André, H.M., Ducarme, X. & Lebrun, P. 2002. Soil biodiversity: myth, reality or conning? Oikos, 96, 3-24. 2. Basset, Y. Horlyck, V. & S.J. Wright (eds) 2003. Studying Forest Canopies from Above: The International Canopy Crane

Network. Smithsonian Tropical Research Institute and UNEP, Panama City. Available on-line at http://www.stri.org/english/research/facilities/terrestrial/cranes/canopy_crane_network.php

3. Basset Y, Mavoungou JF, Mikissa JB, Missa O, Miller SE, Kitching RL & Alonso A. 2004. Discriminatory power of different arthropod data sets for the biological monitoring of anthropogenic disturbance in tropical forests. Biodiversity and Conservation, 13, 709-732.

4. Basset Y, Novotny V, Miller SE & Kitching RL (eds). 2003. Arthropods of Tropical Forests. Spatio-temporal Dynamics and Resource Use in the Canopy. Cambridge University Press.

5. Basset Y, Novotny V, Miller SE, Weiblen GD & Missa O. 2004. Conservation and biological monitoring of tropical forests: the role of parataxonomists. Journal of Applied Ecology, 41, 163-174.

6. Bawa KS, Kress WJ, Nadkarni, NM, Lele S, Raven PH, Janzen DH, Lugo AE, Ashton, PS & Lovejoy TE. 2004. Tropical Ecosystems into the 21st Century. Science, 306, 227-228.

7. Didham RK & Fagan LL. 2003. Project IBISCA – Investigating the biodiversity of soil and canopy arthropods. The Weta, 26, 1-6.

8. Didham RK, Ghazoul J, Stork NE & Davis AJ 1996. Insects in fragmented forests: a functional approach. Trends in Ecology and Evolution, 11, 255–260.

9. Erwin, T.L. 1982. Tropical forests: their richness in Coleoptera and other arthropod species. The Coleopterists Bulletin, 36, 74-75.

10. Hajibabaei M, deWaard JR, Ivanova NV, Ratnasingham S, Dooh RT, Kirk SL, Mackie PM & Hebert PDN. 2005. Critical factors for assembling a high volume of DNA barcodes. Philosophical Transactions of the Royal Society of London, Series B, 360, 1959-1967.

11. Hammond, P.M. 1990. Insect abundance and diversity in the Dumoga-Bone National Park, N. Sulawesi, with special reference to the beetle fauna of lowland rain forest in the Toraut region. In Insects and the Rain Forests of South East Asia (Wallacea), eds W. J. Knight & J.D. Holloway, pp. 197-254. London: The Royal Entomological Society of London.

12. Hammond, P. M. 1995. Magnitude and distribution of biodiversity. In Global Biodiversity Assessment, eds V. T. Heywood & R. T. Watson, pp. 113-138. Cambridge: Cambridge University Press.

13. Hill JK. 1999. Butterfly spatial distribution and habitat requirements in a tropical forest: impacts of selective logging. Journal of Applied Ecology, 36, 364–372.

14. Janzen DH, Hajibabaei M, Burns JM, Hallwachs W, Remigio E & Hebert PDN. 2005. Wedding biodiversity inventory of a large and complex Lepidoptera fauna with DNA barcoding. Philosophical Transactions of the Royal Society of London, Series B, 360, 1835-1845.

15. Kitching RL. 1993. Towards rapid biodiversity assessment – Lessons following studies of arthropods of rainforest canopies. In: Rapid Biodiversity Assessment: Proceedings of the Biodiversity Workshop, Macquarie University – 1993, pp. 26–30. Research Unit for Biodiversity and Bioresources, Macquarie University, Sydney.

16. Lawton JH, Bignell DE, Bolton B, Bloemers GF, Eggleton P, Hammond PM, Hodda M, Holt RD, Larsen TB, Mawdsley NA, Stork NE, Srivastava DS & Watt AD 1998. Biodiversity inventories, indicator taxa and effects of habitat modification in tropical forest. Nature, 391, 72–76.

17. Leponce M, Theunis L, Delabie J & Roisin Y. 2004. Scale dependence of diversity measures in a leaf-litter ant assemblage. Ecography, 27, 253-267

18. Linsenmair KE, Davis AJ, Fiala B & Speight MR (eds). 2001. Tropical Forest Canopies: Ecology and Management. Kluwer Academic Publishers, Dordrecht, The Netherlands.

19. Longino J., Colwell RK & Coddington JA. 2002. The ant fauna of a tropical rainforest: estimating species richness three different ways. Ecology, 83, 689-702.

20. May, R.M. 1990. How many species? Philosophical Transactions of the Royal Society of London, Series B, 330, 293-304. 21. Millennium Ecosystem Assessment. 2005. Ecosystems and Human Well-being: Biodiversity Synthesis. World Resource

Institute, Washington, D.C. 22. Novotny V, Basset Y, Miller SE, Weiblen GD, Bremer B, Cizek L & Drozd P. 2002. Low host specificity of herbivorous

insects in a tropical forest. Nature, 416, 841-844. 23. Pennisi, E. 2005. Sky-high experiments. Science, 309, 1314-1315. 24. Roslin T. 2003. Not so quiet on the high frontier. Trends in Ecology and Evolution, 18, 376-379.

http://www.stri.org/english/research/facilities/terrestrial/cranes/canopy_crane_network.php

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25. Secretariat of the Convention on Biological Diversity. 2000. Sustaining life on Earth. How the Convention on Biological Diversity promotes nature and human well- being. Secretariat of the Convention on Biological Diversity, United Nations Environmental Programme, Montreal.

26. Tangley L. 1990. Cataloging Costa Rica’s diversity. Bioscience, 40, 633–636. 27. UNEP. 2002. Convention on Biological Diversity, Sixth Conference of the Parties. The Hague, April 2002. Document

UNEP/CBD/COP/6/20. Published at www.biodiv.org/ 28. Stork, N.E. 1988. Insect diversity: facts, fiction and speculation. Biological Journal of the Linnean Society, 35, 321-337. 29. Wilson, E.O. 1987. The little things that run the world (the importance and conservation of invertebrates). Conservation

Biology, 1, 344-346. 30. Wilson EO. 2004. Taxonomy as a fundamental discipline. Philosophical Transactions of the Royal Society of London, Series

B, 359, 739.

APPENDIX II. ORGANIZATIONS SUPPORTING IBISCA AND CTRL

Country Organization Contact person(s) Academic institutions

Belgium Royal Belgian Institute of Natural Sciences Dr M. Leponce Université Libre de Bruxelles Prof Y. Roisin & Dr O. Hardy Ministère de la région wallonne Dr M. Dufrêne Czech Republic Czech Academy of Sciences

University of South Bohemia Prof V. Novotny, Dr L. Cizek & M. Janda

Denmark University of Copenhagen Dr L. Sorensen France Université Blaise Pascal Prof B. Corbara Université Toulouse III Prof A. Dejean & Dr J. Orivel CIRAD H.-P. Aberlenc Herbarium, Université de Montpellier II Dr D. Frame Germany University of Erlangen Dr J. Schmidl & J. Bail Universität Würzburg Dr A. Floren United Kingdom The Natural History Museum in London Dr N.D. Springate University of Oxford Prof O. Lewis Zoological Society of London Dr J. Bridle Imperial College S. Pinzon University of Aberdeen Dr O. Missa Roehampton University Prof C. Ozanne Finland University of Helsinki Dr T. Roslin Italy Museo Civico di Storia Naturale, Carmagnola Dr G. Curletti Norway Norwegian Institute for Nature Research Dr F. Ødegaard Switzerland Universität Basel Prof D. Burckhardt Muséum d'Histoire Naturelle de Neuchâtel Dr P. Cuénoud USA National Museum of Natural History in

Washington Dr S.E. Miller

Harvard University Prof E.O. Wilson Lousiana State University Dr A. Tishechkin University of Minnesota Prof G. Weiblen Canada University of Victoria Prof N.N. Winchester Mexico Universidad Nacional Autónoma de México Dr G. Castaño-Meneses

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Panama Smithsonian Tropical Research Institute in Panama Dr D. Roubik, Dr Y. Basset, M. Samaniego & M. Rapp

University of Panama Prof H. Barrios, E. Medianero, A. Barba Brazil Universidade Estadual de Santa Cruz Prof J. Delabie Federal University of Ouro Preto Prof S.P. Ribeiro Universidade Federal de Minas Gerais Dr E. Oliveira Universidade Estadual de Campinas Prof T. Lewinsohn Australia Griffith University Prof R.L. Kitching New Zealand University of Canterbury Prof R.K. Didham Crop & Food Research Dr L. Fagan Research groups United Kingdom Molecular Systematics Section, Royal Botanic

Gardens, Kew Dr V. Savolainen

USA Discover Life Dr J. Pickering Costa Rica Area de Conservación Guanacaste (ACG) Prof D.H. Janzen Arthropods of La Selva (ALAS) Prof J.T. Longino Panama Movimiento de Investigaciones Biológicas de

Panamá A. Cornejo

Papua New Guinea New Guinea Binatang Research Centre (NGBRC) Prof V. Novotny Networks BIOCASE Dr W. Berendsohn Consortium of European Taxonomic Facilities

(CETAF) Dr M. Ramos

European Network for Biodiversity Information (ENBI)

Dr W. Los

EDIT Dr S. Tillier Fauna Europea Dr Y. de Jong Consortium for the Barcode of Life (CBOL) Dr S.E. Miller Convention on Biological Diversity (CBD) H. Zedan Ecoport Dr T. Putter Global Canopy Programme (GCP) A. Mitchell Global Taxonomy Initiative (GTI) Dr R. Hill International Canopy Network (ICAN) Dr N. Nadkarni Pro-Natura International (PNI) O. Pascal United Nations Educational Scientific and Cultural

Organization (UNESCO) N. Ishwaran

United Nations Environmental Programme (UNEP)

Dr M. Oduk



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More information about the support of the IBISCA initiative is available at

http://www.naturalsciences.be/cb/ants/projects/ibisca_main.htm.


United Nations Environment Programme Mission: To provide leadership and encourage partnership in caring for the environment by inspiring, informing, and enabling nations and peoples to improve their quality of life without compromising that of future generations. UNEP has a historical interest in the topic of biodiversity of forest canopies through the IBISCA project. In view of UNEP’s commitment to promote

the scientific base of national decision making of biodiversity and the applications of the ecosystem approach to global/regional biodiversity assessments, it is encouraging to note that the workshop will address the crucial role of the biodiversity of insects, mites and spiders in a biome of global importance i.e. tropical rainforest ecosystem.

UNEP

P.O.Box 30552 Nairobi, Kenya

Ph: +254 2 62 3636 / Fax: +254 2 62 3926 http://www.unep.org

The European Science Foundation (ESF) acts as a catalyst for the development of science by bringing together leading scientists and funding agencies to debate, plan and implement pan-European scientific and science policy initiatives.

ESF is the European association of 78 major national funding agencies devoted to scientific research in 30 countries. It represents all scientific disciplines: physical and engineering sciences, life and environmental sciences, medical sciences, humanities and social sciences. The Foundation assists its Member Organisations in two main ways. It brings scientists together in its EUROCORES (ESF Collaborative Research Programmes), Scientific Forward Looks, Programmes, Networks, Exploratory Workshops and European Research Conferences to work on topics of common concern including Research Infrastructures. It also conducts the joint studies of issues of strategic importance in European science policy. It maintains close relations with other scientific institutions within and outside Europe. By its activities, the ESF adds value by cooperation and coordination across national frontiers and endeavours, offers expert scientific advice on strategic issues, and provides the European forum for science.

European Science Foundation BP 90015

1 quai Lezay Marnésia F-67080 Strasbourg Cedex Fax: +33 (0)3 88 37 05 32

http://www.esf.org

http://www.unep.org/

http://www.esf.org/

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Global Canopy Programme

A global alliance linking studies of forest canopies worldwide into a collaborative programme of research, education and conservation addressing biodiversity, climate change and poverty alleviation.

The aim of the Global Canopy Programme (GCP) is to integrate forest studies across the world into a ten year linked program of research, conservation and education, focussed on understanding the critical role of forest canopies in biodiversity and climate change. It also aims to identify societal benefits from forest canopies, and transmit information to key stakeholders. This initiative evolved from an ESF / NSF funded International Canopy Science Workshop in Oxford, held in November 1999. At the workshop, a template for the GCP was produced by 29 international experts from 10 countries. They concluded that by working together, canopy researchers would be able to leverage more funding for a major collaborative Natural Science project to investigate "nature’s last biotic frontier". They called for significant new funding on the scale of large Physical science projects (US$20-50 million) to undertake this pioneering task.

Global Canopy Programme John Krebs Field Station

University of Oxford Wytham, Oxford OX2 8QJ

United Kingdom Ph : +44 (0) 1865 724 222 ; Fax +44 (0) 1865 724 555

http://www.globalcanopy.org

http://www.globalcanopy.org/

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Convenor: Maurice Leponce [email protected] Tel: +32 2 627 43 58 Fax: +32 2 649 48 25 Co-Convenor: Yves Basset [email protected] Tel: +507 212 8233 Fax: +507 212 8148

Royal Belgian Institute of Natural Sciences Conservation Biology Section 29 rue Vautier 1000 Brussels Belgium Smithsonian Tropical Research Institute Apartado 0843-03092 Balboa, Ancon Panama City Panama

Table of content:

1. Executive summary 2. Scientific content 3. Results and future directions 4. Final programme 5. Abstracts of presentations 6. Final list of participants 7. Statistical information on participants



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1. Executive summary A three-day workshop was organized at the Royal Belgian Institute of Natural Sciences, Brussels, 6-8 July 2005, with financial support from United Nations Environment Programme and the European Science Foundation (www.naturalsciences.be/cb/ants/meetings/esf_exploratory_workshop.htm). The workshop focused on the international project IBISCA (Investigating the BIodiversity of Soil and Canopy Arthopods; public site at www.naturalsciences.be/cb/ants /projects/ibisca_main.htm). This project studies the vertical stratification and beta diversity of arthropods in the San Lorenzo rainforest in Panama, using state-of-the art methods of canopy access and sampling, namely canopy fogging, canopy cranes, single rope techniques and canopy raft and peripherals. The aims of the workshop, which was attended by 37 scientists from 16 countries, were (a) to summarize what has been learned from the IBISCA project overall; (b) to plan meta-data analyses and the dissemination of this novel and important information; and (c) to use this material as a springboard to initiate new collaborative programs of research about the distribution of mega-biodiversity in tropical rainforests and to plan a ‘census of canopy life’. The workshop consisted of 30 presentations and plenary and group discussions. Preliminary results of the IBISCA project. Currently, the IBISCA database which summarizes horizontal (beta diversity), vertical (vertical stratification) and seasonal distribution of arthropods in the San Lorenzo forest includes 85 contributors (ecologists and taxonomists), 14 sampling programmes, 7,233 samples, 422,217 specimens and 1,080 species. We estimate that the final product should shed light on the spatial and seasonal distribution of about half a million of specimens and several thousands of species, distributed among ca 60 focal groups of different phylogeny and ecology. This database has currently no equivalent. The main job of sorting the material to higher taxa and extracting focal taxa has been done for all sampling programmes. The morphotyping of most focal groups will probably be completed by December 2005, with different pace. Identifications, when possible, will be much slower. We expect that databasing of most IBISCA-related information could be completed by June 2006, and that participants will be able at that time to analyze collectively their major findings and report them in a leading scientific journal. Problems identified during the IBISCA project could be eased by working with local parataxonomists trained beforehand to sort and process focal groups, as well as additional funding to speed up different tasks. Analyses and dissemination of IBISCA results. The following steps are needed to complete the IBISCA-Panama project: additional field work, further taxonomical analyses and specimen databasing, processing of ecological variables, improvement of the IBISCA database, and development of a web site. The keys questions targeted by IBISCA are: (1) What is the relative contribution of vertical stratification, seasonality and degree of beta diversity to the distribution of arthropod biodiversity in a closed-canopy tropical rainforest? (2) How do life history traits of species, such as host specificity or feeding guild, influence the spatial and temporal partitioning of

http://www.sciencesnaturelles.be/cb/ants/meetings/esf_exploratory_workshop.htm


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arthropod biodiversity in a closed-canopy tropical rainforest? To this end, one leading concept will be to consider diversity partitioning: total diversity consists of alpha diversity (within sample units), horizontal beta diversity, and vertical beta diversity. So far, the IBISCA project has been presented in 6 scientific articles and 13 magazine articles. We expect that most of IBISCA results should be disseminated during 2006-2008 (multi-authored research papers, high-profile collective article in a leading journal, collective book, etc.). Towards a census of canopy life. The ‘IBISCA’ brand should be retained as it is valuable and should be used in further projects. The IBISCA research group (ca 80 scientists) should also get organized into a non-profit association. The most pressing priority of the organization would be rising funding as to quickly complete the IBISCA-Panama project. Future priorities would be, among others, to persist with the IBISCA-approach beyond the Panama project and to join new biodiversity-related projects, detailed in the text. Improvement of future research needs (a) considering parataxonomist help to facilitate future IBISCA-type projects; (b) focusing on the relations between biodiversity and ecosystem functions, disturbance and climate change; and (c) designing better IBISCA-style projects to help answer geological and evolutionary questions. An ambitious project such as a ‘census of canopy life’ should be implemented in three incremental steps: (1) organize formally the collective group of experts known as ‘IBISCA’ and provide this group with substantial funding; (2) initiate IBISCA-style projects and pilot studies at different locations worldwide and refine the different protocols used in these projects and studies; (3) Develop an aggressive programme aiming at censuing canopy life at key locations in the tropics, based on lessons learn previously. To our knowledge, there are no other alternatives to the above steps if the scientific community is serious about cataloguing canopy arthropods before habitat loss and global climate change extinguish most of this mega-biodiversity.

2. Scientific content The three-day workshop (6-8 July 2005) was attended by 37 scientists from 16 countries. The aims were threefold (hereafter Aims 1-3): 1) to summarize what has been learned from the IBISCA project overall; 2) to plan meta-data analyses and the dissemination of this novel and important information; 3) to use this material as a springboard to initiate new collaborative programs of research about the distribution of mega-biodiversity in tropical rainforests and to plan a ‘census of canopy life’ modeled after the lessons learned from IBISCA. The participants reviewed progresses and impediments of the IBISCA project (Aim 1) during one day and a half, further discussed analyses, dissemination and future projects (Aims 2 & 3) during another day and a half. The workshop consisted of 30 presentations (20 min. each, all available on a CD forwarded to each participant) and plenary and group discussions. Four working groups were established:

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• Gr. 1: IBISCA: a common framework for data analysis: led by Hardy &

Lewinsohn, including Dufrêne, Missa, Leponce, Ribeiro & Roslin. • Gr. 2: IBISCA: taxonomy and interactions between organisms: led by Roisin,

including Aberlenc, Cornejo, Curletti, Delabie, Schmidl, Sorensen, Samaniego & Springate.

• Gr. 3: IBISCA: arthropod distribution patterns: led by Didham, including Bail, Basset, Bito, Floren, Frame, Ødegaard, & Ozanne.

• Gr. 4: Towards a census of canopy life: led by Kitching & Mitchell, including Bridle, Corbara, Cuénoud, Lewis, Fagan, Medianero, Novotny, Oliveira, Pascal & Van Osselaer.

Each working group made recommendations relevant to one or several of the aims of the workshop. In the following section, these recommendations are summarized with regard to the three aims of the workshop.

3. Results and future directions 1. Aim 1: What has been learned from IBISCA? 1.1. Databasing. As of 14 March 2005, the IBISCA database which summarizes horizontal (beta diversity), vertical (vertical stratification) and seasonal distribution of arthropods in the San Lorenzo forest included 85 contributors (ecologists and taxonomists), 14 sampling programmes, 7,233 samples, 422,217 specimens and 1,080 species. We estimate that the final product should shed light on the spatial and seasonal distribution of about half a million of specimens and several thousands of species, distributed among ca 60 focal groups of different phylogeny and ecology. 1.2. Sampling programmes. The IBISCA project consisted of 15 arthropod sampling programme and one vegetation census at all studied sites in the San Lorenzo forest (n=12, one site = 20mx20m). One sampling programme was aborted (butterfly traps) and another one has been re-started in 2005 (rearing of saproxylophagous insects). The results of other sampling programmes are progressively being databased, but with different pace. 1.3. Return of information in the database. So far, Malaise traps, fogging, sticky, light and flight-intercept traps have produced most of the specimens databased, while the number of species databased has been highest for light traps (faster return of information in the database). The vegetation and bee programmes can be considered as being completed, while the results of ground flight-intercept traps and Berlese will be much slower to return to the database, for example. 1.4. Taxonomic analyses. The main job of sorting the material to higher taxa and extracting focal taxa has been done for all sampling programmes. The morphotyping of most focal groups will probably be completed by December 2005, with different pace (example: Apoidea fast, Staphylinidae slow). Identifications, when possible,

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will be much slower. At present, the proportion of morphospecies identified at the species level varies widely (0%: many groups; 25%: Tenebrionidae; 82%: Arctiidae; 100%: Euglossinae and Meliponinae). We expect that databasing of most IBISCA-related information could be completed by June 2006, and that participants will be able at that time to analyze collectively their major findings and report them in a leading scientific journal. 1.5. Positive aspects. The positive aspects of the IBISCA project were in order of decreasing importance (as reflected in the number of opinions voiced by participants): great team spirit, exposure to other colleagues and methods, the integrative nature of the project, and, more distantly, the good logistics of the project. 1.6. Bottlenecks and problems. The most cited impediments were as follows: a) in the field: a1 - pre-sorting of the material could have been better organized, a2 - travel from laboratory to field time-consuming/tedious; b) processing of the material: b1 - lack of time, b2 - lack of funding, b3 - high number of samples and specimens, b4 - label information difficult to match or lost, b5 - quality of insect material low, and b6 - high number of juvenile (inadequate for taxonomy) in the samples; c) taxonomic analyses: c1 - taxonomic knowledge/literature of the Neotropical fauna poor/outdated, c2 - lack of funding to visit collections. 1.7. Remedies to these problems. Problems related to items a1, b1, b3, b4 and b5 above could be eased by working with local parataxonomists trained beforehand to sort and process focal groups. Additional funding would ease problems related to items b2 and c2. Further projects should pay attention to item a2, while items b6 (bar-coding a possible remedy, costs very high) and c1 have no immediate, straightforward solutions. Items 1.8 to 1.11 represent a selection of preliminary results that should be confirmed by rigorous analyses, as agreed by the IBISCA team (item 2.3.1). 1.8. New species. Many species collected during the IBISCA project are new to science and will be described in due course (Oribatida, Milichiidae, etc.). This often includes specimens originating from the upper canopy, but not always. New species may often be cryptic, for example the many species of Anobiidae collected by fogging. 1.9. Beta diversity. Despite the short distance between study sites (< 2km), different sampling methods (Malaise, sticky and flight-intercept traps, fogging, etc.) indicate that arthropod abundance/activity differ greatly between sites. The meaning of this observation is yet obscure, but it may be related to difference in forest productivity, as there appears, for example, to have a positive correlation between arthropod abundance and the Normalized Difference Vegetation Index, as estimated from Landsat pictures (Malaise trap data, Fig. 1). These differences may be stable since

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fogging data suggest that arthropod recolonization after fogging takes place immediately and re-establish a stable rank order and dominance structure.

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Fig.1. Relationship between the average no. of arthropods collected at each site with Malaise traps and forest productivity, as estimated by the Normalized Difference Vegetation index.

1.10. Vertical stratification. Several sampling programmes (light, sticky and flight-intercept traps, beating, notably) suggest rather low faunal overlap along the vertical profile of the forest. This is apparent not only at the species level, but also for higher taxa. For example, the higher taxonomic composition of flight-intercept traps varies much more markedly between vertical heights than horizontally between sites. Arthropod activity follows a bimodal distribution, with greatest activity near the ground and in the upper canopy (flight-intercept and sticky traps, Fig. 2). Patterns of stratification differ among taxonomical and ecological groups, with some ground- or canopy-dominant groups. For example, buprestids of the genus Agrilus, Derbidae, Formicidae, Isoptera, Euglossinae, Oribatida appears more abundant or species-rich near ground, whereas active leaf-galls, Psylloidea, Kalotermitidae, Meliponinae are more abundant/species-rich in the canopy/upper canopy.

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Fig.2. Number of arthropods collected at different heights with sticky (left) and flight-intercept traps (right). 1.11. Seasonality. Fogging data appear to emphasize the important role of yearly variation in rainfall on arthropod abundance at San Lorenzo.

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1.12. Further analyses. Calculating robust statistics to infer sound arthropod distribution is a challenge, since sample size differs among sampling methods, sites and habitats (soil, understrorey, mid-canopy, upper canopy). 2. Aim 2: Analyses and dissemination of IBISCA results 2.1. Completion of the IBISCA-Panama project A certain number of steps are needed to complete the IBISCA-Panama project, so that the questions listed in section 2.2 may be optimally answered. Funds should be raised to this end (item 3.6). 2.1.1. Additional field work. Ideally, the following field studies should be completed in the field in Panama (but perhaps use available data as a guide to simplify field work): (a) measure gradient of disturbance at the sites (e.g., distance to edge, etc.); (b) assess soil typology and slope; (c) complete canopy pin point studies for 4 sites not surveyed during 2003-2004 (Ribeiro). (d) conduct complementary studies (e.g. ant mosaic) 2.1.2. Taxonomical analyses and specimen databasing. Funds should be raised to: (a) speed up the processing and morphotyping of specimens (Oribatida, Hymenoptera Parasitica; ants, etc.); (b) speed up databasing of specimens (e.g. Ødegaard’s beetle focal taxa); (c) visit insect collections (Braconidae: Texas; Curculionoidea+Auchenorrhyncha: InBio in San José, etc.); (d) expand the range of focal taxa (e.g., Collembola). 2.1.3. Processing of ecological variables. All hemispherical digital pictures should be processed to get estimates of canopy openness for the different study sites, and at the different heights sampled (Basset). The acquisition and processing of new remote sensing data (Chust?), with higher resolution pictures than Landsat ones (Ribeiro) may be important to assess forest productivity (Normalized Difference Vegetation Index) at the study sites and possible differences in floral and faunal composition. 2.1.4. Database. A certain number of improvements will be needed for the IBISCA database: (a) determine which plant specimens are located in the understorey/mid-canopy/upper canopy on the basis of regression equations between DBH and height, and calibration in the field (Basset);

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(b) manage better the field ‘height’ (merge some heights) and have a clear definition of ‘habitats’ or ‘strata’ so that we know what are the common heights (strata) across methods; (c) add meteorological and phenological (tree) data in the database (Basset/Wright); (d) propose other ways of expressing arthropod dispersal (e.g. geography, distance) and add such a field in the database; (e) add the fields ‘habitat specificity’ and ‘host specificity’ for each species; (f) measurement of body size and digitized pictures for each morphospecies (item 2.4.3); (g) clean-up of the database (Leponce/Basset); (h) implementation of statistical modules (item 2.3.1). 2.1.5. Guild systems for ecological analysis. Group 2 emphasized that guild systems are essential for structure-, community- and function-related analyses. The extraction of a guild system derived from collected (field) data is better than a theoretical system alone. Schmidl & Springate will prepare a survey sheet and send it to the IBISCA team. Each participant should propose guilds, elements, functions to analyse her/his data appropriately. Schmidl & Springate will compile and implement (after overall agreement from the team) the guild system in the database. Eventually, participants will assign their species/groups to the designated guilds in the database. For comparisons with other studies, the system of arboreal guilds of Moran & Southwood may be retained as a distinct field in the database. 2.1.6. Web site. Ideally, a single IBISCA web site should be implemented (currently two sites are active, one public, one password-protected). This site should include items related to the IBISCA association (item 3.1); the IBISCA database, media reports and scientific articles, etc. Access to different web pages should be either public or password-protected, depending on the sensitivity of the information. As far as possible, the web site should also include digitized pictures of all IBISCA morphospecies (item 2.4.3). It should display a number of important links, especially taxonomic links and funding sources. A person should be hired to develop the interface, manage the web site and the correspondence associated to it (funds needed). 2.2. Analyses of the IBISCA results – questions and hypotheses Group 3 discussed more particularly the questions that IBISCA needs to address from the viewpoint of arthropod distribution (items 2.2.1-2.2.6). 2.2.1. Key questions. Group 3 recommended asking the two following key questions that will optimize the use of information included in the IBISCA dataset: • What is the relative contribution of vertical stratification, seasonality and degree

of beta diversity to the distribution of arthropod biodiversity in a closed-canopy tropical rainforest?

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• How do life history traits of species, such as host specificity or feeding guild, influence the spatial and temporal partitioning of arthropod biodiversity in a closed-canopy tropical rainforest?

2.2.2. Partitioning spatial and temporal components of diversity. See item 2.3.7 (Group 1) for quantifying variance due to beta-diversity and vertical stratification. For the purposes of defining the ‘groups/units’ among which to partition components of diversity, we have 3 major spatio-temporal ‘axes’ of interest: • spatial turnover among sites • vertical stratification • temporal variation among sampling replications We need to quantify these gradients with ecological variables, NOT with arbitrary categories, and we need to identify who is going to lead or help with analyses in these three areas. 2.2.3. Examples. A replication number currently defines temporal variation. It could be better defined by the amount of rainfall during sampling periods (Fig. 3):

1 2 3 4Replication

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Fig.3. Hypothetical relations between arthropod diversity and a gradient of temporal variation.. Another example: vertical stratification: we should abandon height as a surrogate for vertical stratification.. Instead, we should measure the direct biotic and abiotic factors to which species respond (e.g., light, canopy openness, see below). If strata must be delimited, define them as segments of continuous gradients (Didham presentation).

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2.2.4. Potential variables to quantify each gradient: Spatial turnover among sites Differences in floristic composition Plant-related variables, e.g., basal area NDVI (remote sensing) Spatial location / distance between sites Vertical stratification Light (limited data) Canopy openness (crucial to analyze) Leaf area index (Ribeiro - only 5 sites) Temporal variation among Rainfall sampling replications Temperature Tree phenology (flowering, seeding at crane

sites) Hence, arthropod distribution patterns may be predicted from the set of variables measured at each IBISCA site: floristic composition, canopy openness, basal area, etc. 2.2.5. The biological values of singletons and how to deal with them. Although it is obvious that nothing can be said about the life history or biology of singletons (rare species), we should not exclude them from ordination analyses. Ideally, we should: • analyze proportions of singletons in relation to the three ecological gradients; • quantify the reasons why some species appear to be singletons at certain

times/places; • use other data collected on host specificity at the San Lorenzo crane (Ødegaard,

Barrios, Basset, etc.) to interpret why some species are singletons in the IBISCA dataset.

2.2.6. Possible interesting extensions of IBISCA data. The predictive or applied value of IBISCA data could be emphasized to ask the following question: What are the implications for conservation of spatial and temporal partitioning of arthropod communities, with respect to biodiversity loss resulting from logging (canopy openness), land use, or climate change? Data should relate to focal taxa, feeding groups (guild structure), patchiness of forest processes, etc. Group 2 discussed more particularly the questions that IBISCA needs to address from the viewpoint of taxonomy and interactions between species (items 2.2.7-2.2.10).

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2.2.7. Comparison and complementarity of sampling techniques. Group 2 recommended that each participant should screen his/her material for efficiency of methods in the collection of his/her focal taxa, in terms of ecological & taxonomical questions. Ideally, this should be compiled as a chapter in the collective IBISCA book planned (item 2.4.5), with an emphasis on: • selecting focal taxa for different assessments; • selecting methods for sampling these focal taxa; • presenting a manual for optimal sampling protocol(s) in future projects. This chapter should aim at answering how similar are trapping methods and what is their complementarity (could use Chi-square approach in order to compare methods, item 2.3.8). 2.2.8. Interactions between organisms. Group 2 pondered whether it was possible to infer species interactions from distribution patterns in the dataset, by screeening it for correlations of species and guilds. It was thought that, with the exception of a few studies (Ribeiro: galls and plants; Tishechkin: Nitidulidae in ant’s nests; Dejean et al.: ants and plants), interaction data were not measured directly. Since this was not within the aims and scope of IBISCA, the group recommended not to spend too much time on this topic. 2.2.9. Ant mosaics. Group 2 similarly discussed whether ant mosaics exist in the San Lorenzo forest and whether they have an impact on arthropod distribution. This is a complicated search for patterns: e.g. do patches with ant nests support smaller numbers of arthropods (except ants)? It was not clear whether there are enough data and whether it will be possible to extract this information by linking datasets (statistical/mathematical approach). However, given that the majority of IBISCA participant feel that this is an important subject, it may be worth to invest more time on this topic than on item 2.2.8. 2.2.10. Undescribed species. Group 2 was interested to discuss the ratio between described/undescribed species and its bearing on global estimates of biodiversity. These ratios are highly taxon dependent and sound proportions cannot be estimated after thorough taxonomic revisions of the material. If this happens at all, this is unlikely to be within a timeframe of 10 years or more. It was thought that this taxonomic impediment can only be met in part by barcoding, which appears not very realistic or practical for this end. Hence, this topic should not be of high priority to the IBISCA team. In addition to the topics discussed in more details by Groups 2 and 3, there were other items that the IBISCA participants collectively thought were important (items 2.2.11and 2.2.12). 2.2.11. General questions to answer:

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• How to standardise measurements? • Will patterns change based on the chosen standardisation? • How much species turnover is there? • How many species are there in the San Lorenzo forest? • What is the rate of species addition when vertical strata are added to the ground

data? • What is the magnitude of reported patterns for vertical stratification? • What is the top of the canopy? • Is light a surrogate indicator for measuring biodiversity? • Is there a control treatment for IBISCA? • Do we agree on spatial beta diversity across all 9 sites or a nested analysis with

heterogeneity across sites? 2.2.12. Further ideas/questions to consider: • Make an effort to consider questions regarding taxonomic aspects. For example:

write a taxonomic paper to detail how long it takes to identify specimens to a suitable resolution for such a large project and costs involved, etc.

• Do a comparison between insects within guilds to separate the effects of competition (metadata analysis).

• Use the IBISCA taxonomy to build an arthropod phylogeny. Locate the nearest genetic distance for each species from GenBank to produce an overall paper.

• For a general paper that will detail the species richness of focal taxa and turnover among 9 sites: how unique are species to sites, habitats (canopy/understorey) and seasonal components? What are the striking differences among taxa?

• Provide questions on specialisation (canopy vs. ground), processes, herbivory, edges, climate.

2.3. Analyses of the IBISCA results – statistical considerations 2.3.1. Working group 1. This group will implement a common set of statistical analyses (on the IBISCA web site) that will allow the IBISCA group to report collectively their results in a leading scientific journal around mid-2006. The recommendations of working groups 2 and 3 will be used to perform key analyses for this article. Group 1 should communicate with other IBISCA participants via the IBISCA web site. Ideally, funding should be raised so that this group could meet very soon in Brussels, where the majority of group participants reside. 2.3.2. Species accumulation curves. They will constitute the basis of many analyses and should be computed across taxa, sites and habitats. 2.3.3. Steps for analyses. Group 1 suggested: 1. Defining first the basic units of sampling (4 dimensions: sampling methods; habitats (understorey/upper canopy); time (replications); and space: horizontal, study sites).

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2. To consider the description of the sampling (its intensity and ecological variables associated), in order to appreciate the ecological distribution (e.g. linked to plant distribution) vs. spatial distribution. 3. To ask what is the biodiversity of the San Lorenzo forest and use descriptive statistics of sampling results. 4. To consider the sampling efficiency and evaluate methods (or combination of methods) for future projects. 5. To characterize diversity patterns. 6. To explain species assemblages. 2.3.4. Example. For a summary paper on diversity patterns in canopy vs. lower layers, Group 1 recommended: 1. To reduce the complexity of the sampling. This would include to analyze separately 1 focal taxon collected with one sampling technique, keep one season (September-October 2003), and keep only data relevant to canopy and lower layer (understorey, ground). However, this suggestion was not well received by all participants, some fearing loss of information. 2. To describe the samples, using a raw characterization of the number of species and individuals, per taxon-technique-stratum. 3. To perform statistical analyzes, including diversity comparisons, diversity partitioning, and comparison of horizontal and vertical turnover (items 2.3.5-2.3.7). 2.3.5. Diversity comparisons. First, calculate rarefaction curves for all samples per basic unit, canopy vs. lower layer (comparison per sampling unit and/or individual). Second, compare Fisher’s alpha, and Simpson’ diversity. Procedure: extract the data from database and use the software EstimateS. 2.3.6. Diversity partitioning. Either the species richness or Simpson’s diversity could be partitioned between two factors: vertical effects (canopy vs. lower) and horizontal effects (5-9 study sites). Thus, total diversity may consist of alpha diversity (within sample units), horizontal beta diversity, and vertical beta diversity (Fig. 4). The interaction between horizontal and vertical beta diversity is of special interest. Procedure: check within software PRIMER.

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Alpha

(within sampling unit)

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Fig.4. Partitioning of species richness or diversity with relevance to IBISCA data.. 2.3.7. Comparison of horizontal and vertical turnover. Consider the pairwise similarity coefficient between sampling units [Morisita-Horn (= NESS(1) or NESS (100?)]. Plot this similarity vs. the spatial distance (or log (distance)) (Fig. 5).

3 curves: Within canopyWithin lower layer

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Fig.5. Plot of faunal similarity against distance for different situations. Procedures: Morisita-Horn is calculated with EstimateS, NESS (100) with COMPAH96. Regressions and Mantel tests can be calculated in Excel and PC-ORD. ANOVAs on the intercept a and slope b may be calculated for different guilds (are there general trends?). 2.3.8. Chi-square analyses. Didham and Fagan also suggested considering a simple test: comparing the observed frequency of catches of a particular taxon at different forest levels with the expected catches of that taxon (based on total catches of all arthropods at different forest levels) with a Chi-square test (Fig. 6).

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Fig.6. Chi-squared test of the occurrence of dolichopodid catches at various height above the ground.. 2.3.9. Ideas/questions to consider. These include notably: • how to analyze distances between sites (nested, combined, other); • which beta diversity calculation will be used and reported collectively; • should we attempt to analyze data for phylogeny and independent contrasts: if

possible yes and look at different insect traits: body size, distribution, life history, dispersal, etc;

• the independence of data points and how to test for it; • meta–data style analyses allowing consideration of all sampling methods and

specimens collected during IBISCA: standardize taxa abundance by reporting the relative abundance per sample, not sums;

• what is the best way to visually present our data?

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2.4. Dissemination of IBISCA results 2.4.1. Present dissemination and expected output. So far, the IBISCA project has been presented in 6 scientific articles and 13 magazine articles (see recent pubs in section V). A number of manuscripts have been initiated. We expect that most of IBISCA results should be disseminated during 2006-2008. 2.4.2. New species and deposition of material. Group 2 discussed this topic and it was agreed that Sorensen will prepare guidelines to be sent to taxonomists together with IBISCA material: • deposition of holotypes • reference collection for Panama • copyright issues? • selling names to fund IBISCA taxonomists? 2.4.3. Virtual reference collection and web site. Group 2 and the majority of IBISCA participants thought that a IBISCA web site should be maintained (item 2.1.6) and that it should also include an internet catalogue of IBISCA morphospecies/species as a virtual reference collection. Ideally, an export routine from the database into html files should be written. It would display data / pictures (upper/underside, details) / descripions / references / literature, etc., as well as (important) information on the collection where the material is deposited. If possible, look for existing models and collaborate with other groups, (Binatang Research Center, ALAS). 2.4.4. Strategies for dissemination. Collectively, IBISCA participants should use the following strategies to disseminate IBISCA results (in order of decreasing importance): • Multi-authored research papers in peer-reviewed scientific journals. • High-profile collective article in a leading journal. • Collective book, equivalent to the proceedings of the present workshop. • Publications in scientific magazines of wider scope such as Trends in Ecology

and Evolution or Scientific American. • IBISCA web site (items 2.1.6, 2.4.3). 2.4.5. Research papers. The general strategy to publish research papers would be as follows: 1) Draw up a list of potential papers. Participants outline and submit paper ideas to Basset, who list them on the IBISCA web site. 2) Each participant to highlight which paper they want to be included as an author. The authorship will be finalised by all working on the paper. Any disagreement could result in authorship being alphabetized.

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3) As far as possible, funds should be raised to cover page fees (if any) for the main IBISCA articles. 2.4.5. High-profile collective article. Hopefully, this collective article could be drafted towards mid-2006. Basset suggested that the authorship of this paper should refer as ‘IBISCA’, with an appendix listing all participants (including taxonomists) involved in the article (perhaps 50-60 authors). There have been precedents, such as papers published by ‘APG’ (Angiosperm Phylogeny Group). This would help to retain the brand name ‘IBISCA’ for future projects (item 3.1). 2.4.5. Collective book. IBISCA has so far raised ca $15,000 towards editing a collective book free of copyright on the preliminary results from the IBISCA-Panama project. The project will be innovative as providing in one package: • First rate science: preliminary results of IBISCA and reports originating from the

present workshop. Perhaps 40 multi-authored chapters of 10 pages each, plus references and index.

• Attractive iconography: many colour pictures, perhaps organized in plates (2-3 plates per chapter if possible).

• Searchable companion CD. • Public access to downloadable pdf files and additional information (pictures,

updated species lists), hosted on the IBISCA web site. • Free dissemination to targeted readers and groups (scientists and institutions in

developing countries: university librairies, IBISCA participants): perhaps 500 copies.

• Nominal fee for other readers: perhaps 500 copies at $5-10 each, to cover mailing expenses associated with the above item.

The final product should be aimed as a cross between a standard edited scientific book and a richly illustrated ‘coffee table’ book. This represents one of the best ways to promote the study of canopy tropical biology. Further funding should be raised as to secure first rate iconography from professional photographers who participated to IBISCA fieldwork, book handling and mailing expenses, and the creation of html files and CDs. Publication could be achieved towards the end of 2006. Basset will compare the different publishing options (Bulletin RBINS, Panama/Colombia, BioForm, Pensoft, etc.) and prepare an example of chapter. 3. Aim 3: Toward a census of canopy life 3.1. The IBISCA ‘brand’. Group 4 advised to retain the ‘IBISCA’ brand as it is valuable (the exact acronymic meaning is unimportant) and should be used in further projects. The advantages of this strategy are related to: • the whole-forest focus; • the scale of scientific collaboration (which is not exclusive);

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• the fact that it is not just a census (integrative approach); • the future: same approach but different questions. Subsequent discussions among IBISCA participants in the meetings of Leipzig & Uberlandia in July 2005 also established the need for the IBISCA research group (ca 80 scientists) to get organized into a non-profit association, or some similar sort of formal organization. The exact procedure will be explored by Basset, Corbara, Barrios & Leponce, but is likely to include secretarial costs, including the implementation of the new organization. 3.2. Present priorities. Group 4 also listed the following items as present priorities for IBISCA: • Complete the IBISCA-Panama project (section 2.1) and a high profile manuscript

(item 2.4.5) by July 2006, if possible. • Scan the IBISCA data for inferences & hypotheses about disturbance, in order to

secure a potential integrating theme for a future project. • Consider ecosystem functioning. This is rather challenging but our guild analyses

have the potential to be better than others and could be of great help. • Consider the integration of IBISCA data with other datasets. However, the

comparability or integration of data may be more an issue for a future analysis/strategy.

• Raise funding as to quickly complete the IBISCA-Panama project and, in particular, taxonomic analyses (item 3.7).

3.3. Future priorities. Group 4 listed the following items as future priorities for IBISCA: • Persist with the IBISCA-approach beyond the Panama project • The primary (& saleable) issues for future IBISCA-style projects should be the

following key organising principles: disturbance, climate change and ecosystem function.

• The experimental approach is powerful and could be pursued by IBISCA-style projects, thus adding value to existing plots/project. For example, the Kyoto plots, Danum biodiversity experiment, Critical Size of Ecosystem projects come to mind.

• Address evolutionary questions and preserve specimens to allow molecular work • Include medical entomologists in future projects, since disease/vector issues are

related to disturbance and climate change (connection with human welfare). • Consider the opportunity to join new biodiversity-related projects (item 3.5). 3.4. Perspectives and improvement of future research. The majority of IBISCA participants agreed on the following points, which were further discussed by Group 4:

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• Follow-up projects in Panama would be welcome, as they would be supported by existing and mounting information related to the San Lorenzo forest.

• Work on a broader geographic scale is also advisable. • It is crucial to consider parataxonomist help to facilitate future IBISCA-type

projects. It could be project-related or developed as an international ‘parataxonomy facility’.

• IBISCA may need smaller working groups to develop future work plans, for example: ecosystem function / disturbance /climate; refining inventory methods; and future projects.

• Example: the future BATH project calls for process-related research ideas. • Design better IBISCA-style projects to help answer geological and evolutionary

questions. • Sign a declaration for taxonomic sharing in canopy research in all the major

countries involved and send this declaration to each government. • Consider the use of a social scientist for the future IBISCA projects to help avoid

conflicts, confrontations, framework for group dynamics, etc. 3.5. New projects. In addition to new projects in Panama taking advantage of the mounting information on the San Lorenzo forest, new projects keen to involve the IBISCA model may be initiated at the following locations: • Australia: ‘BATH’ project (Kitching, 2006-2007): connection with climate

change. • Brazil, Malaysia, India, Madagascar, Ghana: ‘Whole Forest Observatories’

(Global Canopy Programme/GEF, 2006-2011): develop baseline surveys. • Vanuatu: project SANTO (Corbara, 2006-2007). • French Guiana: Tree glider (Corbara, 2007). • Panama (ANAM/GEF, 2008?). However, this will involve shifting emphasis on ecosystem functioning (i.e., studying processes from ground to upper canopy) and a greater use of the parataxonomist taskforce, as developed by Novotny & Missa in New Guinea and Gabon, for example. 3.6. Donations and fund-raising. During the workshop, one observer, C. Van Osselaer – Biotrac sprl, made a donation of 1,000 € to IBISCA, in order to encourage further activities of the team. IBISCA should raise further funds to: • complete the IBISCA-Panama project and in particular to speed up taxonomical

activities, the processing of ecological variable and improving the database; • disseminate the results of the above project; • explore parataxonomist training in the context of large projects such as IBISCA; • do a pilot study in Panama: expanding canopy access points from crane

perimeters; • do a pilot study in Australia, related to BATH project: ecosystem functions;

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• do a pilot study in French Guiana with the Tree Glider: mobile canopy access; • do a pilot study for a follow-up of IBISCA in Panama (ANAM-GEF); • cover costs for the IBISCA web site and the organization of the association; A detailed budget, probably in the order of 300-500K € is has been prepared by Basset and will be circulated to all participants for comments. Biotrac sprl or the Patron of IBISCA, Prof. E.O. Wilson, could help identifying potential donors. Alternatively, these activities could be sponsored by a joint application of multiple European laboratories to ESF (‘À la carte Scientific Programmes’). OVERALL CONCLUSIONS IBISCA started as a relatively modest project in Panama, with limited participants, funding and opportunities to extend it as to cover a substantial part of arthropod diversity and address the effects of seasonality. Due to the great team spirit and the enthusiasm of participants, IBISCA later grew into a full-scale international project with a core funding of ca $300,000. In the field, IBISCA has demonstrated the value of an integrated project to survey tropical arthropods and has provided answers as to how to tackle the study of this mega-diversity efficiently. The lively intellectual exchanges witnessed during the present workshop promise that the IBISCA team will complete the Panamanian project and disseminate the important mounting information on the arthropods of the San Lorenzo forest. An ambitious project such as a ‘census of canopy life’, which essentially includes collecting countless unknown arboreal tropical arthropods, should be implemented in three incremental steps: 1) Organize formally the collective group of experts (ca 80) known as ‘IBISCA’ and provide this group with substantial funding. 2) Initiate IBISCA-style projects and pilot studies at different locations worldwide and refine the different protocols used in these projects and studies. 3) Develop an aggressive programme aiming at censuing canopy life at key locations in the tropics, based on lessons learn previously. To our knowledge, there are no other alternatives to the above steps if the scientific community is serious about cataloguing canopy arthropods before habitat loss and global climate change extinguish most of this mega-biodiversity. RECENT IBISCA PUBLICATIONS Cízek, L. & Hauck, D. 2005. Jeráby v pralese. Vesmir, 84, 38-43. [in Czech] Corbara, B., Basset, Y. & Barrios, H. 2005. IBISCA: a large-scale study of arthropod

mega-diversity in a neotropical rainforest. In: Tropical Biodiversity: Science, Data, Conservation. Abstract Volume. 3rd GBIF Science Symposium, 18-19

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April 2005, Brussels, Belgium (ed by H. Segers), pp. 31-32. Belgian Clearing-House to the Convention on Biological Diversity, Royal Belgian Institute of Natural Sciences & Global Biodiversity Information Facility, Brussels.

Ribeiro, S.P. & Corbara, B. 2005. Dossel florestal: a fronteira desconhecida. Ciencia Hoje, 217, 54-58.

Roisin, Y., Dejean, A., Corbara, B., Orivel, J. & Leponce, M. 2005. Arthropod biodiversity in tropical rainforest canopies: Panamanian termites in the framework of the IBISCA project. In: Tropical Biodiversity: Science, Data, Conservation. Abstract Volume. 3rd GBIF Science Symposium, 18-19 April 2005, Brussels, Belgium (ed by H. Segers), p. 32. Belgian Clearing-House to the Convention on Biological Diversity, Royal Belgian Institute of Natural Sciences & Global Biodiversity Information Facility, Brussels.

Schmidl, J. & Corbara, B. 2005. IBISCA - Artenvielfalt der Boden- und Baumkronen-Arthropoden in einem tropischen Regenwald (San Lorenzo NP, Panama). Entomologische Zeitschrift, Stuttgart, 115, 104-108.

Pennisi, E. 2005. Sky-high experiments. Science, 309, 1314-1315 A full list of IBISCA publications may be downloaded at http://www.naturalsciences.be/cb/ants/projects/ibisca_reprints.htm

4. Final programme

The workshop was preceded by a public exhibition in the Parc du Cinquantenaire / Jubelpark in Brussels (23-29 June 2005). The exhibition presented the Solvin-Bretzel (“canopy raft”) and informed the public about arthropod biodiversity in tropical forests and the IBISCA project. It was organized by the Royal Belgian Institute of Natural Sciences (RBINS) with the support of Solvay (www.sciencesnaturelles.be/cb/ants/exhibition/ibisca.htm). The posters of the exhibition were presented for a further week at the RBINS (1-8 July 2005).

The canopy raft and the IBISCA exhibition in Parc du Cinquantenaire, Brussels.

http://www.naturalsciences.be/cb/ants/projects/ibisca_reprints.htm

http://www.sciencesnaturelles.be/cb/ants/exhibition/ibisca.htm

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The three-day workshop (6-8 July 2005) started with a pre-workshop welcome day where participants visited the RBINS exhibition and were greeted by Dr Camille Pisani (Director, RBINS), Robert Van Geyts (communication manager, Solvay) and Dr Bruno Corbara (Co-PI, IBISCA project). Dr Corbara presented the IBISCA approach and his talk was followed by a short video, “Mission: IBISCA” (Solvay, 2004).

Tuesday 5 July 2005

Afternoon Arrival

Welcome day

17:30 Welcome of participants and visit of the IBISCA exhibition (sponsored by Solvay S.A.) at the Royal Belgian Institute of Natural Sciences (Museum, 6th level, insect hall).

19:00 Welcome. The involvement of the Royal Belgian Institute of Natural Sciences in biodiversity studies Camille Pisani, general director of the Museum

19:05 The involvement of Solvay in the IBISCA project Robert Van Geyts, communication manager, Solvay

19:10 Presentation of the IBISCA approach Bruno Corbara, scientific director of the Canopy Raft Consortium

followed by a movie (DVD IBISCA) (Main auditorium, Museum)

19:30 video "Mission: IBISCA" (Solvay, 2004) (Main auditorium, Museum)

20:30 Dinner at the restaurant Kapolino

Wednesday 6 July 2005

Distribution of tropical mega-biodiversity: available data

1. Introduction and background 08:30 Presentation of the European Science Foundation (ESF)

Lucien Hoffmann (Standing Committee for the Life, Earth and Environmental Sciences)

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08:50 Introduction to and aims of the workshop (Leponce & Basset)

09:00 Presentation of workshop participants (led by Basset & Leponce)

09:15 Presentation of the IBISCA project: aims and general methods (Corbara)

09:30 IBISCA at a glance: results of the participant survey (Basset & Leponce)

2. Beta diversity

09:40 Beta diversity: basic concepts and tropical data Novotny

10:00 Beta-Diversity: Contributions from ant studies to the debate Delabie

10:20

Coffee Break

10:40 The IBISCA Malaise trap programme: focal taxa and non-formicid hymenoptera. Springate, Basset & Pinzón

11:00 The IBISCA canopy fogging programme and team focal taxa Schmidl, Floren & Bail.

11:20 The IBISCA Winkler programme and focal taxa Aberlenc, Leponce, Orivel, Corbara & Roisin

11:40 The pitfall trap, ground flight-intercept trap, vegetation programmes plus focal taxa: Histeridae, Nitidulidae, Pselaphinae, Ceratocanthidae and Braconidae. Medianero, Tishechkin, Samaniego, Hernandez, Cuenoud, Ribeiro, Barrios & Basset

12:00

Lunch

3. Vertical gradients 13:00 Overview of theory and current concepts of arthropod vertical

stratification Didham

13:20 Higher Questions: unfinished business with canopy arthropods Kitching

13:40 Vertical stratification of beetles in dead branches (The IBISCA beating programme) Ødegaard

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14:00 Moth assemblages. Target groups: Geometridae, Arctiidae, Pyraloidea

(The IBISCA light trap programme) Kitching, Oliviera, Basset & Cornejo

14:20

The IBISCA flight-intercept trap programme (FL) and focal taxa (Psocoptera, Diptera) Fagan, Didham, Rapp, Cuénoud, Swann

14:40 The IBISCA sticky trap programme and focal taxa: Auchenorrhyncha and Agrilus (Hemiptera, Coleoptera) Basset, Curletti, Barrios, Cizek, Aberlenc, Leponce & Barba

15:00

Coffee Break

15:20 Herbivory rates and gall-forming species and density distribution in the canopies of Neotropical ecosystems: from savannas, semi-deciduous forests to wet rainforests. Ribeiro & Vieira

15:40 The microarthropod, beating and wood-rearing programmes plus focal taxa: Oribatida, Tenebrionidae, Clavicornia, Isopoda, Myriapoda & Curculionoidea (minus Scolytinae). Winchester, Jordan, Cizek, Barrios, Ødegaard Curletti & Basset

4. Interactions between organisms

16:00 Trophic interactions among tropical organisms Lewis

16:20 The IBISCA social insects programme: ants Orivel, Leponce, Delabie, Corbara, Roisin, Cardoso do Nascimento, Ribeiro, Seniuk, Esteves, Campos, Samaniego, Jordan, Winchester, Schmidl, Floren & Dejean.

16:40 Distribution of termites from the ground to the canopy of a Panamanian rainforest Roisin, Dejean, Corbara, Orivel & Leponce

17:00 Euglossine and meliponine bees diversity and abundance on the ground and in the canopy. Frame & Roubik

5. Integration for the sessions of the day 17:20 Beta diversity (led by Delabie & Novotny)

17:40 Vertical gradients (led by Didham & Kitching)

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18:00 Tropical interactions (led by Lewis & Orivel)

20:00 Dinner at “Stekerlapatte”, in Marolles neighbourhoud

Thursday 7 July 2005

Survey and analyses of mega-biodiversity distribution in the tropics: pitfalls and remedies

08:30 Evolution along selective gradients: linking population genetics with

community ecology Bridle

08:50 Implementing large-scale surveys and experiments in the tropics Roslin

09:20 Taxonomic impediment Sørensen

09:40 Working with parataxonomists Missa

10:10

Coffee Break

10:30 The IBISCA database Leponce & Basset

10:50 Characterizing community diversity in species rich systems – statistical pitfalls, remedies, and insights from a phylogenetic perspective Hardy

11:10 Meta data and multivariate analyses of large datasets Dufrêne

11:30 Whole forest observatories: An international network for monitoring canopy biodiversity and global climate change Mitchell

11:50

Lunch

13:00 Plenary preparation of working group 1: Problems and remedies

related to the analyses of IBISCA data (led by Hardy & Lewinsohn)

13:40 Plenary preparation of working group 2: Key analyses of IBISCA data: taxonomic aspects and interactions between organisms (led by Roisin)

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14:20 Plenary preparation of working group 3: Key analyses of IBISCA data: arthropod distribution patterns (led by Didham)

15:00

Coffee Break

15:30 Plenary preparation of working group 4: Towards a census of canopy life (led by Kitching & Mitchell)

20:00

Dinner at “Les Salons d’Atalaïde”

Friday 8 July 2005 Discussion, integration and conclusions

8:30-12:00

Working group 1: Problems and remedies related to the analyses of IBISCA data (led by Hardy & Lewinsohn, including Dufrêne, Missa, Leponce, Ribeiro & Roslin)

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8:30-12:00

Working group 2: Key analyses of IBISCA data: taxonomic aspects and interactions between organisms (led by Roisin, including Aberlenc, Cornejo, Curletti, Delabie, Schmidl, Sorensen, Samaniego & Springate)

8:30-12:00

Working group 3: Key analyses of IBISCA data: arthropod distribution patterns (led by Didham, including Bail, Basset, Bito Floren, Frame, Ødegaard & Ozanne)

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8:30-12:00 Working group 4: Towards a census of canopy life (led by Kitching

& Mitchell, including Bridle, Corbara, Cuénoud, Gama de Oliveira, Lewis, Fagan, Medianeiro, Novotny, Pascal & Van Osselaer)

10:00-10:20

Break for all working groups

12:00

Lunch

13:00 Working groups 1-4: preparation of a summary of the discussions

14:15 Report of working group 1 (led by Hardy & Lewinsohn)

14:50 Report of working group 2 (led by Roisin)

15:25

Coffee Break

15:45 Report of working group 3 (led by Didham)

16:20 Report of working group 4 (led by Kitching & Mitchell)

16:55 How to ensure an efficient dissemination of IBISCA data (led by Corbara & Leponce) [includes discussion about the workshop proceedings; perhaps formation of ‘writing groups’, discussion of CD, web site, etc.]

17:30 Intervention of participants who did not give a talk

18:00 Conclusions (led by Kitching)

19:00 Beer at Le Roy d’Espagne, Grand Place.

20:00 Picture group in front of Manneken pis.

20:30 Dinner at “Le Manneken”, Grand Place

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5. Abstracts of presentations All presentations are available online (password protected): http://www.naturalsciences.be/cb/ants/meetings/EW04-049-presentations.htm Distribution of tropical mega-biodiversity: available data

Beta diversity Beta diversity: basic concepts and tropical data V. Novotny Department of Ecology and Conservation Biology Institute of Entomology Branisovska 31, CZ 370 05 Ceske Budejovice, Czech Republic The paper reviews main causes of species turnover in space, including speciation, dispersal limitation, habitat availability and biotic interactions. Further, it outlines the basic approaches to measuring beta diversity, relying on pair-wise comparisons between communities or local-regional comparisons. Finally, the paper reviews the little we know about beta diversity of insects in tropical rainforests and discusses promising directions for the further study. Beta-Diversity: Contributions from ant studies to the debate J. H. C. Delabie Laboratorio de Mirmecologia, Centro de Pesquisas do Cacau, CEPLAC, & Departamento de Ciências Agrarias e Ambientais, Universidade Estadual de Santa Cruz, Ilheus, Bahia, Brazil. Currently: Invited Professor, Laboratoire d’Evolution et Diversité Biologique, Université Paul Sabatier, Toulouse, France. According the classical definition of MacArthur, the Beta Diversity is the component of total diversity that can be attributed to differences in species composition among the homogeneous units in the landscape [1]. Ants and other social insects are specially interesting for this kind of study in tropical habitats [2, 3]. Data of the ant communities of southern Bahia studied during the last 15 years for several purposes are used here for considerations about the Beta Diversity partition of ants in tropical latitudes and compared with data obtained through similar sampling efforts in a temperate region. Several topics have to be considered for ant communities studies, according the scale and distribution of the samplings, the vegetation strata, as well as the links with its natural complement concepts that are the alpha- and gamma-diversities. Ants are seen between the dominant organisms of tropical ecosystems. In tropical arborous eco/agrosystems, their communities are radically differently structured in the canopy and on/in the ground. In the former, they are strongly organized in mosaics around dominants and co-dominants species, while this kind of organization, even it exists sometimes in the latter, is much more discreet and can be seasonal. Another important element of the strata differences, consequences of the ant evolutionary history [4], is that their dynamics of reproduction and colonization, self-organization and resource uses are absolutely different, which makes hard a generalization of the Beta Diversity concepts using simultaneously hypogaeic, epigaeic and arborous ant assemblage data. Other studies focus the species turnover according a distance gradient in tropical and temperate homogenous habitats, as well as the community organization in function of the land degradation in a tropical region. Simple models are suggested for the Beta Diversity according the temperate/tropical and anthropization gradients. Finally, as ant ecologists have recently beneficed of the generalization of the Winkler trap method and ant collects standardization [5], some developments of the methods currently applied are suggested, aiming to reach comparative data of the Beta Diversity for further developments of ant communities’ studies.

http://www.naturalsciences.be/cb/ants/meetings/EW04-049-presentations.htm

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[1] Gering, J.C. & Crist, T.O. 2002. Ecology Letters 5: 433-444; [2] Schroeder, J.H. et al. 2004. J. Biogeography 31: 1219-1226; [3] Galbiati et al. 2005. Sociobiology 45: 925-936; [4] Wilson, E.O. & Hölldobler, B. 2005. P.N.A.S. 102: 7411-7414; [5] Agosti, D. et al. (orgs) Ants: Standart Methods for Measuring and Monitoring Biodiversity, Smithsonian Institution. The IBISCA Malaise trap programme: focal taxa and non-formicid hymenoptera. N. Springate, Y. Basset & S. Pinzón In February, 2004, 9 Malaise traps were emplaced in the San Lorenzo Protected Area in ‘plots’ in which plant diversity had been surveyed. The intentions were two-fold: (i) to provide additional material from the field-layer for the surveys of IBISCA focal taxa and (ii) to test possible correlations between the taxonomic and lifeway diversity of non-formicid Hymenoptera and that of angiosperms. Raw data and some provisional analyses, based on abundance of the focal taxa, are presented and discussed. Within rapid biodiversity assessment the need for an alternative approach to the analysis of taxa which occur at low abundance but at high species richness is discussed in the context of non-formicid Hymenoptera. The IBISCA canopy fogging programme and team focal taxa J. Schmidl1, A. Floren2, J. Bail1. 1Ecology & Nature Conservation, Institute for Zoology I, Staudtstr. 5, D-91058 Erlangen, Germany. 2Lehrstuhl für Tierökologie und Tropenbiologie, Universität Würzburg, Biozentrum, Am Hubland, D-97074 Würzburg, Germany. Fogging programme: A systematic canopy fogging programme was conducted within the IBISCA-project, sampling eight sites (F1,F2,F3,B1,B2,I1,R1,R3, each with 6 sub-samples of 5x5m plastic sheets installed in 1m height) in October 2003 and re-fogging six sites (F1,F2,F3,B2,I1,R3) in May and October 2004, using the identical location of the initial fogging. We used a Swingfog SN1 and a 1% natural Pyrethrum solution, dissolved in white oil Essobayol 82. A total of approx. 79000 arthropods was collected by canopy fogging, in average 3942 per site and 657 per 20sqm-sampling-sheet. The table below gives the total numbers for the replications, sites and main insect orders and taxa. Due to the general interest, the Formicidae are listed separately. The total number will be up to 5% (estimated) higher, as the Blattodea and the Residuals were not counted in each season. Arachnida are not counted and separated properly, but Araneae form the main fraction, followed by Opilionids, Pseudoscorpions, Ricinulei and Scorpions. Exact numbers will be available after data input in the database. The rank order of the taxa and its totals is marked greyish. With each re-fogging the number of arthropods sampled in each sub-sample was rising. It is unlikely that this a seasonal phenomenon, as the results for October 2003 and October 2004 show. It highlights the low long-term impact of Pyrethrum both on arthropod communities and ecosystem. A re-colonisation takes places immediately, and the general bionomics represented by different higher arthropod taxa re-establish a +- stable rank order and dominance structure.

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Table: Overview results of canopy fogging October 2003, May 2004 and October 2004, with numbers for each replication and main insect orders/taxa. Ranking of main taxa and underlying total numbers marked greyish. nc: not counted.

October 2003: F123 B12 I1 R13 8 sites 48 sheets

May2004: F123 B2 I1 R3 6 sites 36 sheets

October 2004: F123 B2 I1 R3 6 sites 36 sheets

total 120 sheets / 20sqm

Formicidae 9226 Formicidae 8783 Diptera 9681 24578 Diptera 4532 Diptera 5922 Formicidae 6569 20135 Coleoptera 4425 Hymenoptera 2981 Hymenoptera 2893 9781 Hymenoptera 2854 Coleoptera 2802 Coleoptera 2554 8728 Arachnida 1232 Arachnida 1308 Homoptera 1739 4132 Homoptera 832 Homoptera 712 Orthopteroidea 630 3283 Orthopteroidea 892 Orthopteroidea 789 Arachnida 1592 2311 Heteroptera 313 Heteroptera 478 Heteroptera 380 1171 Thysanoptera 81 Thysanoptera 151 Thysanoptera 130 362 Isoptera 12 Isoptera 3 Isoptera 30 45 Residual 1669 Residual nc Residual nc 1669 Blattodea nc Blattodea 706 Blattodea 444 1150 n total 26080 24635 28117 78832 n per sheet 543 684 781

Focal taxa covered by “Fogging-team”: JSchmidl: ARXXXX, BLXXXX, OPXXXX, COADER, COANOB, COANTC, COARTE, COBOST ,COBYRR ,COCANT, COCERO, COCHEL, COCIID, COCLER, COCOLY, CODASC, CODERM, CODRYO, COELAT, COELMI, COEUCI, COEUCN, COHETE, COLAGR, COLAMP, COLIMN, COLISS, COLYCI, COLYME, COMELA, COMELO, COMELY, COMONO, COMORD, COMYCE, COMYCT, CONOSO, COOEDE, CORHIC, CORHIP, COSALP, COSCIR, COSCRA, (COTENE), COTHRO, COTROG, COZOPH; AFloren: COCHRY; JBail: MAMANT, ORACRI, OREUMA, ORGRYA, ORGRYL, ORGRYT, ORPROS, ORPYRG, ORTETR, ORTETT, ORTRID. The IBISCA Winkler programme and focal taxa H.P. Aberlenc, M. Leponce, J. Orivel, B. Corbara & Y. Roisin The pitfall trap, ground flight-intercept trap, vegetation programmes plus focal taxa: Histeridae, Nitidulidae, Pselaphinae, Ceratocanthidae and Braconidae. E. Medianero, A. Tishechkin, M. Samaniego, Hernandez, Ph. Cuenoud, S. Ribeiro, H. Barrios & Y. Basset Vertical gradients

Overview of theory and current concepts of arthropod vertical stratification R. Didham University of Canterbury, Christchurch, New Zealand In this talk I will give an overview of central concepts in the study of vertical stratification in forests, and how these may relate to the analysis and interpretation of data from project IBISCA and future ‘mega-biodiversity’ initiatives. Although there has been contentious debate about whether true stratification in forest structure exists at all, it is now generally accepted that non-uniform patterns in the abundance and diversity of organisms across vertical heights do occur at some times and places in most forests. Exactly how to define these patterns of stratification, and how to interpret their ecological and evolutionary significance, is another matter altogether. This

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talk will be phrased as a series of discussion questions, rather than as a definitive statement of current knowledge, reflecting the poorly-defined state of the subject. First, I will comment on the forest structural templet on which arthropod vertical stratification is overlaid, and then discuss how best to define stratification and what the key determinants of vertical stratification might be. Subsequently, I will present a series of discussion questions (and caveats) on how to measure, test and interpret vertical trends in the IBISCA data, with particular consideration of the appropriate null model(s) we should be using to test for the existence (and magnitude) of vertical stratification. Finally, I will comment on the relevance of vertical stratification to the global distribution of biodiversity, with the conclusion that the importance of vertical stratification cannot be judged independently from the relative magnitude of beta diversity across vertical heights. Higher Questions: unfinished business with canopy arthropods R. Kitching Griffith University There are many exciting areas of ecology and environmental science in which canopies and canopy arthropods play vital roles. This talk presents a personal view of some exciting questions for the post-IBISCA era. The evaluation of 'good' canopy science rests on four pillars: connecting pattern with process, connecting data with theory, pursuing achievable goals, and demonstrating societal relevance without compromising scientific quality. Decomposition processes in the canopy are intriguing. Studies of perched litter in Asplenium allow a range of exciting questions to be pursued - relating to biodiversity patterns, biodiversity/process connections and the testing of 'geand' ecological theories. Recent ground-breaking work on herbivory leads directly(like all good science)to more questions. Ready canopy access and potential collaborations with plant eco-physiologists allow us to target local scale heterogeneity in the herbivory process. Scaling up from herbivory studies, there are huge opportunities to do food-web work based on guild analyses in forest canopies. Community webs in microhabitats and both source and sink webs in the broader canopy environment can be constructed. Forest to forest comparisons using the forest observatory network will be rewarding. Perhaps the biggest challenge of all for students of canopy arthropods is quantifying the connections between arthropod-driven processes in the canopy and local climate. The connection is indicated in recent work on volatile organic carbons. Mid-length carbon compounds, VOC's, are involved as cloud seeds over tropical forests - maintaining forest quality and ameliorating climate change. In forests VOC's are biogenic products produced in response to plant-animal interactions. Following either forest clearance or direct anthropogenic VOC production, these compounds combine with nitrogen oxides to release ozone - exacerbating global warming. We need to know the biologically connections involved. A network of canopy observatories will permit key observations and manipulations to quantify these connections. Vertical stratification of beetles in dead branches (The IBISCA beating programme) F. Ødegaard The aim of this side project of IBISCA was to compare the vertical stratification of saproxylic beetles within and between different tree species. The study was preformed during October 2003 and May 2004 at the Canopy Crane site in the San Lorenzo forest in Panama. For each of 18 different tree species, four freshly cut branches were suspended in the canopy (15 to 25m above ground) and placed in the understorey (1m above ground) of their parent tree, respectively. All branches were beaten regularly (approximately every third day) for four weeks in both sampling periods. There were a total of 10 bouts of beating on each bunch of branches. All Coleoptera associated with dead wood and senescing leaves were collected. The study yielded ca. 4,937 beetles belonging to ca. 661 species. The results showed that both abundance and species richness

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was significantly higher in the canopy than in the understorey. The canopy fauna was more host specific than the understorey fauna. Species richness, abundance and host specificity did not change across seasons. Detrended correspondence analyses showed that the beetle fauna within strata (canopy and understorey) of different tree species were more similar than between strata of conspesific tree species. A summary of the status of sorting, identification and databasing of my focal taxa is presented below. This includes all material received by 9. June 2005. The rest of the unsorted material, which includes about 5-10 000 individuals, I expect to receive soon. Group Species Individuals Identified % id. % db Unsorted? Carabidae 88 398 34 38.6 100 100 Scydmaenidae 85 492 3 3.5 100 100 Scaraboeoidea 86 632 74 86.0 100 1000 Bruchidae 9 17 6 66.7 100 10 Hispinae 19 35 13 68.4 100 10 Chlamysinae 3 4 0 0.0 100 0 Cerambycidae 124 410 89 71.8 100 50 Anthribidae 32 71 9 28.1 100 50 Attelabidae 8 93 6 75.0 100 10 Scolytinae 193 12000 82 42.5 5 5000 Platypodidae 20 261 18 90.0 100 100 There are no big problems in the sorting and identification process, but several factors contribute to slow down the progress. 1) Lack of funding for sorting. 2) Lack of time for handling such huge material 3) I haven’t received the last parcels that includes about 25% of the total material. I expect data to be ready for final analyses earliest by end of 2005 and latest by end of April 2006. Moth assemblages. Target groups: Geometridae, Arctiidae, Pyraloidea (The IBISCA light trap programme) R. Kitching1, E. Oliviera2, Y. Basset3, A. Cornejo4 1Griffith University, 2 University of Ouro Preto, 3Smithsonian Tropical Research Institute, 4University of Panama As of the end of June, a total of 5246 specimens sampled by light traps had been added to the data base. Of these 2185 were Lepidoptera and, of these 1784 belonged to the four target families. There were 421 arctiids, 385 geometrids, 774 crambids and 204 pyralids (s.s.). In addition 493 non-target Lepidoptera have been processed but, generally, not identified. No doubt many more data from Orders other than Lepidoptera await attention. With the aid of the NMNH collections in Washington, 63% of target taxa have been identified to species, 78% to genera. Comparable percentages for each family are: Arctiidae - 82 & 94, Geometridae - 78 & 97, Crambidae - 57 & 75, Pyralidae - 32 & 38. The pyralid subfamilies Phycitinae, Acentrominae (=Nymphulinae) and Epipaschiinae proved most intractable for identification. Clear vertical patterns exist in the data even though more formal analysis is yet to be done. The Geometridae, for example, show a clear pattern at the level of the subfamily with sterrhines dominating the ground fauna, ennomines the canopy. Formal statistical analysis can now be done involving ordination and Estimates approaches.

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The IBISCA flight-intercept trap programme (FL) and focal taxa (Psocoptera, Diptera) L. Fagan1, R. Didham2, M. Rapp3, P. Cuénoud4, J. Swann5 1Crop & Food Research, Lincoln, New Zealand 2Canterbury University, Christchurch, New Zealand 3Canterbury University, Christchurch, New Zealand 4Conservatoire et Jardin botaniques dela Ville de Genève, Genève, Switzerland 5University of British Columbia, Vancouver, Canada Ordinal abundance patterns from 1,659 flight-intercept trap (FL) samples for all IBISCA focal taxa are briefly summarized. A total of 71, 721 specimens were captured in FL traps at six heights from the forest floor to the canopy, from September 2003 to October 2004. Coleoptera, Hymenoptera and Diptera represented 75% of the FL catch. Proportional representation of taxa was relatively constant between sites, but varied markedly between vertical heights. In particular, Coleoptera increased two-fold in abundance and proportional representation from the ground to the canopy. Focal Coleoptera families will be the most important taxa to analyse from FL samples. With respect to focal Psocoptera from FL traps, there was also a strong increase in abundance an species richness with increasing vertical height. For focal Diptera taxa across all IBISCA trapping programmes (N=190,461), we present a brief breakdown of progress on sample sorting, focal families to be sorted to morphospecies and results obtained so far. There were strong patterns of vertical stratification at the family-level for all Diptera and at the genus/species-level for Milichiidae (Diptera). Apparent canopy-dominant families include Dolichopodidae, Chloropidae, Milichiidae and Scatopsidae. Ground-dominant families include Cecidomyiidae, Drosophilidae, Phoridae, Sphaeroceridae. For Milichiidae, species composition is extremely unusual for a Neotropical assemblage, and canopy-level sampling has forced a re-evaluation of major sampling methods the are useful for collecting Milichiidae, and of the relative dominance of the genera Phyllomyza and Pholeomyia in the Neotropics. The IBISCA sticky trap programme and focal taxa: Auchenorrhyncha and Agrilus (Hemiptera, Coleoptera) Y. Basset1, G. Curletti2, H. Barrios3, L. Cizek4, H.-P. Aberlenc5, M. Leponce6, A. Barba7 1Smithsonian Tropical Research Institute, Panama, PA. 2Museum of Carmagnola, Carmagnola, IT 3University of Panama, Panama, PA 4Czech Academy of Sciences, Ceske Budejovice, CZ 5CIRAD, Montpellier, FR 6Royal Belgian Institute of Natural Sciences, Brussels, BE 7University of Panama, Panama, PA This contribution summarizes progress with the IBISCA sticky trap programme and presents preliminary results for two focal taxa: Auchenorrhyncha and Agrilus (Buprestidae). In total, the sticky trap programme surveyed 993 traps at 9 sites and yielded ca. 55,000 arthropods. Arthropod abundance/activity along the vertical transect follows a bimodal distribution and is significantly higher (and of similar magnitude) at the levels of soil/litter and upper canopy. Patterns of vertical stratification greatly differ among arthropod groups with different ecologies. Incident light measured below the traps appears to be a good predictor of the abundance/activity of arthropods collected per trap. About 15,000 homopterans (Auchenorrhyncha and Psylloidea) representing 446 morphospecies were collected with a variety of sampling methods during the IBISCA project. About 72% and 29% of this material was identified at the generic and species levels, respectively. Taxonomical studies are on-going. Stratification and faunal turnover is obvious at familial, subfamilial and specific levels. More species were collected in the understorey (where sampling effort was highest), but rarefaction curves were similar for the understorey and the upper canopy, with the mid-canopy being enriched from both habitats. Adults whose nymphs are fungal/root feeders are prevalent near the forest ground, whereas meristem-feeders dominate in the upper canopy. The distance (perhaps related to floristic composition) and the illumination of the sites

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appear important to predict homopteran species richness at each site, not numbers of plant species per se. Extreme specialists appear uncommon and specialization appears to occur more towards vertical distribution than site. Fifty-eight species of Agrilus are known from Panama. Most Agrilus gathered during IBISCA (n = 56 specimens) were collected with sticky traps. This material included 19 species, out of which 12 are new for Science and will be described shortly. More individuals and species were collected in the understorey, especially in forest gaps, than in the upper canopy. We estimate that at least 58 species of Agrilus must occur in the San Lorenzo forest. These results emphasize that (a) the Agrilus fauna is very poorly known in Panama; (b) that this fauna appears to be richer in the understorey than in the upper canopy; and (c) that tree-fall gaps are important for the maintenance of local diversity. Herbivory rates and gall-forming species and density distribution in the canopies of Neotropical ecosystems: from savannas, semi-deciduous forests to wet rainforests. S. Ribeiro & A. Vieira Universidade Federal de Ouro Preto The present protocol explores canopy versus understorey vegetation and habitat structure, and herbivory/galling insect data. All galls were counted and measured, and data entry for vegetation sample is finished. 2004 data needs analyses, but general figures are already available. Some plant data won’t be trivial to analyse, and causa mortis for galls are not easy to identify. Final analyses may be finish by August/September. Using the “canopy/understorey cylinder” method, a specific forest volume could be quantified and compared. Understorey had in average 7.3 times more plant individuals than the canopy, but the canopy presented 8.5 times more leaves in this same volume, in average for all studied sites. Site B1 understorey had the greatest leaf area index (nearly double than the average, and 6 times greater than C3, the least dense understorey). Similarly, the leaf area index for B1 canopy was the greatest, being 2.3 times greater than the average, and 2.7 times greater than the raft site, the least dense canopy. Data support the hypothesis that gall-forming insect population distribution and survivorship is highly correlated to sampling height within the forest: there are more galls in high branches (multiple linear regression, F2,27=6.9, p < 0.02), which also had the most sclerophyllous leaves in the whole forest (simple linear regression, F1,28=7.09, p < 0.01). Galls in the understorey had greater mortality rates than in the canopy. Nevertheless, high infestation was detected only on saplings of canopy tree species. Host tree effect is important but needs further analyses at this stage. Sclerophylly has been proposed as an important mechanism in favour of gall-forming survivorship, and the present data comes in support of this hypothesis. In addition, sclerophylly could prevent free-feeding herbivores (chewing) activity, which could also be an auxiliary mechanism in favour of gall forming oviposition site choice, based on finding harsh habitats, where sclerophyllous leaves will prevail. Accordingly, herbivory rates were significantly higher in the understorey plants, consistently across all sites, and regardless the significantly smaller amount of resources in this forest habitat (ANOVA mixed model, F5,237=3.0, p<0.01). Furthermore, herbivory rates decreased significantly in the canopy with sample height (simple linear regression, F1,247=33.4, p < 0.001). The present work may change the perception of ecophysiological patterns along canopy vertical gradients, and the proper methods to study such habitat. Within-plant traits may be as much or more relevant for insect herbivore distribution than micro-climatic conditions. In addition, data calls for more studies on the highly specialists endophagous insects. The microarthropod, beating and wood-rearing programmes plus focal taxa: Oribatida, Tenebrionidae, Clavicornia, Isopoda, Myriapoda & Curculionoidea (minus Scolytinae). N. Winchester, K. Jordan, L. Cizek, H. Barrios, F. Ødegaard G. Curletti & Y. Basset

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Interactions between organisms

Trophic interactions among tropical organisms O. Lewis Institution (s) University of Oxford Tropical insect herbivores, their host plants and their predators and parasitoids account for the vast majority of the earth's biodiversity. Studying trophic (feeding) interactions among these species in diverse tropical ecosystems creates special challenges, but has the potential to further our understanding of the processes structuring and maintaining patterns of diversity and abundance. I will briefly review approaches used to study plant-herbivore and herbivore-parasitoid interactions in tropical forests, and describe the use of food webs to quantify interactions across multiple trophic levels. Such studies allow us, for the first time, to make robust and testable predictions about the implications of adding or removing species from ecological communities. I will consider how IBISCA data might contribute to ongoing work on trophic interactions, and how data on tropical insects might in future be collected to maximise its value to the study of trophic interactions. The IBISCA social insects programme: ants J. Orivel, M. Leponce, J.H.C. Delabie, B. Corbara, Y. Roisin, I. Cardoso do Nascimento, S. Ribeiro, Seniuk, Esteves, R. Campos, M. Samaniego, K. Jordan, N. Winchester, J. Schmidl, A. Floren & A. Dejean. Distribution of termites from the ground to the canopy of a Panamanian rainforest Y. Roisin1, A. Dejean2, B. Corbara3, J. Orivel2, M. Leponce4 1Behavioural and Evolutionary Ecology, CP 160/12, Université Libre de Bruxelles, Avenue F.D. Roosevelt 50, B-1050 Brussels, Belgium. 2Laboratoire d'Evolution et Diversité Biologique, UMR-CNRS 5174, Bât. 4R3, Université Toulouse III, 118 route de Narbonne, F-31062 Toulouse cedex 4, France 3LAPSCO, UMR-CNRS 6024, Université Blaise Pascal, 34 avenue Carnot, F-63037 Clermont-Ferrand cedex, France 4 Section of Conservation Biology, Royal Belgian Institute of Natural Sciences, Rue Vautier 29, B-1000 Brussels, Belgium Termites are inhabitants of warm temperate or tropical ecosystems. Numerous studies focusing on fallen logs, leaf litter or humus have established their importance as decomposers at ground level, but almost no attention has been paid to their presence in the upper strata of tropical forests. Within the framework of the IBISCA project in the San Lorenzo Protected Area (Panama), we conducted the first systematic sampling campaign to evaluate the diversity and richness of a termite fauna, from the ground to the canopy. Dead wood or termite-built covered runways were examined on a total of 125 trees along two transects, whereas quadrats provided samples of the ground fauna at the same sites, for comparative purposes. Canopy collections (here defined as higher than 10 m above ground) yielded 63 occurrences (colony samples) representing 10 termite species, whereas 29 species were recorded in 243 occurrences from the ground. Five species were recorded in both habitats. Species accumulation curves revealed that the inventory of canopy species was near completion, whereas ground species were still accumulating in a logarithmic pattern. Remarkable components of the canopy fauna include several drywood species (Kalotermitidae), forming small colonies within dead branches or stumps. By contrast, soil feeders were exclusively found in ground samples, where they were abundant (19 species, 110 occurrences). Wood feeders displayed a similar species richness at both levels, although most species showed a clear preference for either ground or canopy. Further data still to be analyzed include: 5 ground transects (> 350 series) in other sites of the same forest, to evaluate Beta-diversity; 27 series of additional hand-collected canopy samples; 78 series of non-flying termites collected by standardized sampling methods (Berlese, Winkler, pitfalls, etc.); and > 370 samples of flying termites (alates) collected by light or flight interception traps, yet to be identified. Put

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together, those data should provide an integrated picture of the termite fauna of this forest, including vertical and horizontal richness and distribution patterns. Euglossine and meliponine bees diversity and abundance on the ground and in the canopy. D. Frame & D.Roubik STRI and Herbarium, Insitut de Botanique, Univ. de Montpellier II As part of the IBISCA project, we surveyed euglossine and melipoinine bee diversity and abundance on the ground and in the canopy. I will present a description of the protocol we used to sample these bees and our sampling results. The data was analyzed using a similarity index, the so-called Morsita Horn Index. The aim of this presentation is to provide a basis for discussion of how to best integrate the bee results into the overall IBISCA framework and data analysis. Survey and analyses of mega-biodiversity distribution in the tropics: pitfalls and remedies Evolution along selective gradients: linking population genetics with community ecology J. Bridle Institute of Zoology, ZSL, London NW1 4RY A central question in population genetics concerns what limits evolutionary responses to ecological change. Traditionally this issue has been considered in terms of adaptation to a single selective dimension, based on a balance between natural selection and migration between populations along a spatial gradient. In the long term, the way that populations can track changing conditions in time determines the generation and maintenance of diversity (speciation), and therefore the degree of specialisation and complexity observed in natural communities. I will discuss some of the evolutionary factors they may constrain the niche width of species, and discuss how the selective gradient might be extended to measure the selective gradient in terms of ecological interactions between and within species, in addition to the effects of gene flow within species. Implementing large-scale surveys and experiments in the tropics T. Roslin Metapopulation Research Group,Department of Biological and Environmental Sciences PO Box 65 (Viikinkaari 1),FI-00014 University of Helsinki, Finland How can we identify the processes behind the megadiversity that this workshop focuses on? In my talk, I will try to identify some broad types of approaches to experiments and surveys of arthropod communities in the tropical forest. As a population biologist, I will focus less on studies on mere species richness, and more on studies attempting to combine information on individual species and their habits with patterns of species diversity. And as a temperate biologist, I will borrow freely from studies conducted by others. In particular, I will examine the role of ‘model systems’, as compared to a more even focus on a larger set of species and/or sites. Overall, I will claim that flagship projects such as IBISCA are urgently needed to increase the public appeal, credibility and competitiveness of biodiversity research, but that results from single locations must eventually be validated by studies conducted at a broader range of sites. I will also contend that as far as the circumstances permit, work at the community level should be linked to an assessment of species-specific ecology and population-level processes. I will try to illustrate the (partial) feasibility of (some of) these claims by a recent project on the dung beetles of Madagascar. Taxonomic impediment L. Sørensen The Natural History Museum of Denmark , University of Copenhagen, Denmark

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The taxonomic impediment comprises two components, one spatial and one temporal. The first of these is that the majority of the world’s biodiversity is confined to the tropics, while most taxonomic capacity is found in the “developed” world. The second is that taxonomic capacity is decreasing, while the need for taxonomic information is increasing. This taxonomic impediment has stimulated ongoing discussions and initiatives to amelioriate the problem, such as digitizing museum collections, DNA “bar coding” of species, biodiversity megadatabases. How will these initiatives help when taxonomic capacity is so limited and knowledge of fauna and flora so incomplete for most of the biodiverse tropical countries. Will we have to wait until all species are described? What are the ways forward and are there any shortcuts? This paper gives an overview of possible strategies and tools to reduce the problems arising from the taxonomic impediment Working with parataxonomists O. Missa Parataxonomists are typically local people with no formal education in biology, who stand "at the side" of professional taxonomists and biologists and help them in the acquisition of biological information. In the past, the parataxonomist's role has often been limited to helping biologists collect samples in the field. More recently, however, parataxonomist activities have expanded to include sorting (at a variety of levels from families to morpho-species), databasing, preparing specimen and even digital imaging. Given the proper training and feedback there are few repetitive tasks that parataxonomists could not perform reliably for professional biologists. Although involving parataxonomists in a biodiversity inventory can be very productive, it is also paved with potential pitfalls. A strategy must therefore be put in place to guarantee that data quality is high and remains constant throughout a project. I conclude this short talk by presenting my own personal views on how to involve parataxonomists with maximum efficiency in a project like IBISCA. The IBISCA database M. Leponce & Y. Basset The IBISCA database currently contents more than 50,000 records including 400,000 specimens classified into 2,189 taxa (species or higher level). Taxa already identified up to species level are principally: Coleoptera (Anthribidae, Carabidae, Cerambycidae, Chrysomelidae, Curculionidae, Scarabaeidae, Scydmaenidae), Hemiptera (Achilidae, Cicadellidae, Cixiidae, Delphacidae, Derbidae, Flatidae, Issidae, Membracidae, Psyllidae), Hymenoptera (Apidae, Formicidae), Isoptera and Lepidoptera (Arctiidae, Geometridae, Pyralidae). Characterizing community diversity in species rich systems – statistical pitfalls, remedies, and insights from a phylogenetic perspective. O. Hardy Eco-éthologie Evolutive. Université Libre de Bruxelles. Assessing species diversity of rich communities is a difficult task due to the numerous sources of biases that can occur when collecting data and when analyzing them. I will overview the common pitfalls and discuss some remedies, essentially from a statistical point of view (which estimators to choose). Some links between diversity coefficients and neutral community models will be mentioned. I will show the potential interest of partitioning diversity coefficients into alpha and beta components (within versus among sites, sampling units, local habitats,…) for inferential purposes. I will also illustrate the potential insights that can be obtained by integrating phylogenetic information into the analysis of community structure. Finally, I will try to demonstrate that all this can be overviewed in 15 minutes. Meta data and multivariate analyses of large datasets M. Dufrêne Centre de Recherche de la Nature, des Forêts et du Bois, Ministère de la Région wallonne, Gembloux

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Broad species inventories produce huge datasets with many information on species presence or frequencies in numerous samples. The identification of main biological structures or ecological relationships can be then quite difficult. In such a situation multivariate analyses aim 1) to summarize main biological or ecological structures or 2) to reveal relationship between ecological factors and biological structure. All data matrix where rows are samples and columns are biological descriptor (as species abundance), ecological factor (as environmental factor values) or metadata (as species number or life history features) can be visualized in a graphical way, using columns as coordinate axes where samples are projected in function of their column values. All samples form then a cloud of point-objects that can have peculiar structure, revealing strong gradients (when descriptors are well correlated) or aggregated sub-clouds or clusters (when only several samples share the same descriptor combination) or a combination of gradients and clusters or no structure at all. To reveal main gradients, the sample cloud is swivelled on its barycentre to be viewed to maximize its larger lengths (= its variance) : samples are then projected in a new coordinate system with the first main axes pointing to the larger lengths and where these axes are simply a linear combination of the original descriptors. On these main axes, the rank or the ordination of samples allow to identify those that are the most different and located at the extremes of the gradients. Classical descriptive methods for extracting main gradients are Principal Components Analysis (matrix of ecological descriptors, linear relationship), Correspondence Analysis (matrix of species frequencies, unimodal distribution) and Principal Coordinate Analysis (based on any metric distance matrix). For cluster approach, a large diversity of similarity indices and cluster methods (hierarchical or not) exist and it is important to choose those corresponding to descriptor properties (value distribution, ordinal or not, ...). Gradient and cluster analyses are really two complementary ways to describe a data matrix structure because the first one privileges great distances between samples then the second reveals strong similarities among samples. The interpretation of biological or ecological structure revealed by ordination analyses can be done respectively with simple correlation between the sample rank on main axes and other independent descriptors that have not been included on the ordination analysis. By example, one can revealed the main biological gradient opposing samples on the basis of species frequencies with a CA on a samples/species abundance dataset (what are the samples that are the most different when we use a species abundance list ?). Such true species gradient can be after correlated with several ecological descriptors or their combination (multiple regression) to identify what could be the responsible main limiting ecological factors or the ecological factors that explain the ecological niche partitioning. For sample clusters, simple classical Anova (one independent factor at the same time) or better discriminant analyses (to identify a combination of independent factors) are often sufficient. One classical example is the identification of ecological factors that explains sample clusters obtained after the realization of an UPGMA or Ward clustering method on a Steinhaus or Bray-Curtis similarity matrix computed on a samples/species abundance dataset. A peculiar case of sample cluster interpretation is the identification of indicator species, i.e. species that can be almost systematically associated to groups of samples. Twinspan and IndVal approaches are two methods largely used to solve this problem. In all the cases listed above, the aim is to explain one depend variable (the position on a gradient or the membership to a sample group) by one or a combination of explaining factors. Such approaches are generally called indirect (gradient or cluster) analyses; the main biological or ecological structure are identified and after, one searches to interpret it. Recent developments of different techniques allow now a direct (gradient) analyses, i.e. the explanation of a collection of depend variables by a combination of explaining factors. One of the most known technique is the Canonical Correspondence Analysis where the CA axes (privileging unimodal response between species and ecological factors) obtained on a samples/species abundance dataset are constrained and modified to be also the better linear combination of available ecological factors. In such case, the main axes of the sample cloud are not defined by the most extreme species but by the most extreme species that can be explained by a linear combination of available ecological factors. The

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quality of the ecological response is maximized but it depends on the quality of the species dataset and also, on the quality of the ecological factor dataset. Similar approaches exist for linear relationship between dependent and independent variables (Redundancy Analysis) or between similarity matrix (Mantel test) or between dendrograms (consensus indices). RA is by example an interesting way to identify relationship between several expressions of species number (guilds, families, life history features, ...) and ecological variables describing the environment of samples. A key characteristic of such approaches is the possibility to measure the relation between two datasets independently of a third one (i.e. partial relation ), to control by example spatial autocorrelation or a sampling structure (covariable dataset). In some case, this will allow a quite complete partitioning of the dataset variance (% explained by some kind of factors, by spatial location, by sampling structure, ...). In spite of their strong capacity to clarify the biological and ecological structures, these multivariate methods depend crucially on the quality of the experimental protocols and thus of the formulation of the starting assumptions. The power of the methods is expressed as well as possible only when the questions are clear and that the experimental protocols or of inventories were conceived to answer it positively or negatively. Such approaches cannot correct datasets obtained with protocols not well structured. Too often, the ratio between number of samples and explaining factors is far to be sufficient. It should be greater than 3 and better when there are10 times more samples than explaining factors. Whole forest observatories: An international network for monitoring canopy biodiversity and global climate change A. Mitchell

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6. Final list of Participants Convenor: 1. Maurice LEPONCE Conservation Biology Section Royal Belgian Institute of Natural Sciences 29 rue Vautier 1000 Brussels Belgium Tel: +32 26 27 43 58 Fax: +32 26 49 48 25 Email: [email protected] Co-Convenor: 2. Yves BASSET Smithsonian Tropical Research Institute, Apartado 2072 Balboa Ancon Panama City Panama Tel: +507 212 82 33 Fax: +507 212 81 48 Email: [email protected] ESF Representative: 3. Lucien HOFFMANN Cellule de Recherche en Environnement et Biotechnologies Centre de Recherche Public-Gabriel Lippmann 41, rue du Brill L-4422 Belvaux, Grand-duchy of Luxembourg Tel: +352 470261-400 Fax: +352 470264 E-mail : [email protected] UNEP Representative: Margaret M. ODUK Biodiversity and Biotechnology Unit Division of Environmental Conventions United Nations Environment Programme P.O Box 30552, Nairobi. Kenya Tel: 254-20-623465 Fax: 254-20-624255 Email: [email protected]

Staff: Conservation Biology Section Royal Belgian Institute of Natural Sciences 29 rue Vautier 1000 Brussels Belgium Isabelle BACHY Tel: +32 26 27 43 63 Email: [email protected] Thibaut DELSINNE Tel: +32 26 27 43 64 Email: [email protected] Chris KERWYN Tel: +32 26 27 43 54 Email: [email protected] Yves LAURENT Tel: +32 26 27 43 59 Email: [email protected] Participants: 4. Henri-Pierre ABERLENC CIRAD TA 40/L Campus International de Baillarguet (CSIRO) 34398 Montpellier Cedex 5 France Tel: +33 467 59 31 23 Fax: +33 467 59 31 01 Email: [email protected] 5. Johannes BAIL Ecology & Nature Conservation Department Institute for Zoology I University of Erlangen-Nuremberg Staudtstr. 5 91058 Erlangen Germany Tel: +49 9131 8528076 Fax: +49 911 5188572 Email: [email protected] 6. Darren BITO The New Guinea Binatang Research Center P. O. Box 604











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Madang Papua New Guinea Tel: +675 852 1587 Fax: +675 852 1587 Email: [email protected] 7. Jon BRIDLE Zoological Society of London Institute of Zoology Regent's Park London NW1 4RY United Kingdom Tel: + 44 207 449 6334 Fax: + 44 207 586 2870 Email: [email protected] 8. Bruno CORBARA LAPSCO-UMR CNRS 6024 Université Blaise Pascal 34 avenue Carnot 63037 Clermont-Ferrand Cedex France Tel: +33 4 73 40 62 53 Fax: +33 4 73 40 64 82 Email: [email protected] 9. Aydeé CORNEJO Movimiento de Investigaciones Biológicas de Panamá (MIBio) Ave. Justo Arosemena y Ave. Cuba, Edif. Poli, piso 10, local 10A22. Tel: 262-2305 E-mail: [email protected] ; [email protected] 10. Philippe CUENOUD Conservatoire et Jardin botaniques de la Ville de Genève 1, ch. de l'Impératrice Case postale 60 1292 Chambésy/Genève Switzerland Tel: +41 22 418 51 00 Fax: +41 22 418 51 01 Email: [email protected] 11. Gianfranco CURLETTI Museo Civico di Storia Naturale Cascina Vigna Cas. Post. 89 10022 Carmagnola TO Italy Tel: +39 0119724390 Fax: +39 0119724237 Email: [email protected]

12. Jacques DELABIE Université Toulouse 3 118 route de Narbonne 31062 Toulouse cedex France Tel: :+33 5 61 47 44 95 Fax: +33 5 61 55 73 27 Email: [email protected] 13. Raphael DIDHAM Zoology Department University of Canterbury Private Bag 4800 Christchurch New Zealand Tel: +64 3 364 2059 Fax: +64 3 364 2024 Email: [email protected] 14. Marc DUFRENE Ministere de la region wallonne Centre de recherche de la nature, des forets et du bois Avenue Maréchal Juin 23 5030 Gembloux Belgium Tel: +32 81 62 64 29 Fax: +32 81 61 57 27 Email: [email protected] 15. Laura FAGAN Landcare Research Department PO Box 69 Gerald Street Lincoln New Zealand Tel: +64 3 325-6701 ext. 3815 Fax: + 64 3 325-2418 Email: [email protected] 16. Andreas FLOREN Biozentrum Lehrstuhl Zoologie III Universität Würzburg Am Hubland 97074 Würzburg Germany Tel: +49 931 8884376 Fax: +49 931 8884352 Email: [email protected] 17. Dawn Maria FRAME Laboratoire de Botanique Tropicale Institut de Botanique Université de Montpellier II















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163, rue A. Broussonet 34090 Montpellier France Tel: +33 4 67577180 Fax: +33 4 67570180 Email: [email protected] 18. Evandro GAMA DE OLIVEIRA Conservation International, TEAM Research Scientist Tropical Ecology, Assessment and Monitoring (TEAM) Initiative Brazil Tel: +55 31 3498-6444 Email: [email protected] 19. Olivier HARDY Laboratoire d'Eco-éthologie Evolutive Université Libre de Bruxelles CP 160/12 Ave F.D. Roosevelt 50 1050 Bruxelles Belgium Tel: +32 2 650 6585 Fax: +32 2 650 2445 Email: [email protected] 20. Roger KITCHING Research Centre for Tropical Rainforest Ecology and Management Australian School of Environmental Studies Griffith University Nathan Brisbane QLD 4111 Australia Tel: +61 738 757 491 Fax: +61 738 755 014 Email: [email protected] 21. Thomas LEWINSOHN Laboratório de Interações Insetos-Plantas Depto Zoologia Instituto de Biología Universidade Estadual de Campinas CP 6109 13083-970 São Paulo Brazil Email: [email protected] 22. Owen LEWIS Department of Zoology University of Oxford South Parks Road Oxford OX1 3PS United Kingdom

Tel: +44 1865 271162 Fax: +44 1865 310447 Email: [email protected] 23. Enrique MEDIANERO Programa de Maestría en Entomología University of Panama Panama City Panama Tel: +507 264 54 31 Fax: +507 264 54 31 Email: [email protected] 24. Olivier MISSA Zoology Building School of Biological Sciences University of Aberdeen Tillydrone Avenue Aberdeen AB24 2TZ United Kingdom Tel: +44 1224 27 2393 Email: [email protected] 25. Andrew MITCHELL Global Canopy Programme Department of Zoology University of Oxford John Krebs Field Station Wytham Oxford OX2 8QJ United Kingdom Tel: +44 1865 724 222 Fax: +44 1865 724 555 Email: [email protected] 26. Vojtech NOVOTNY Czech Academy of Sciences Dept of Ecology and Conservation Biology Institute of Entomology University of South Bohemia Branisovska 31 37005 Ceske Budejovice Czech Republic Tel: +420 38 530 0350 Fax: +420 38 530 0354 Email: [email protected] 27. Frode ØDEGAARD Norwegian Institute for Nature Research Tungasletta 2 7485 Trondheim, Norway Tel: +47 73 80 15 55 Fax: +47 73 80 14 01 Email: [email protected]














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28. Claire OZANNE Centre for Research in Ecology & the Environment School of Human & Life Sciences Roehampton University Holybourne Avenue London SW15 4JD United Kingdom Tel: +44 20 8392 3598 Email: [email protected] 29. Olivier PASCAL Pro Natura International 15, avenue de Ségur 75007 Paris France Tel: +33 1 53 59 97 98 Fax: +33 1 53 59 94 46 Email: [email protected] 30. Sérvio RIBEIRO Departamento de Biologia Geral/ICB Federal University of Ouro Preto, Universidade Federal de Minas Gerais CP 486 30161-970 Belo Horizonte MG Brazil Tel: +55 3135591665 Fax: +55 3135591660 Email: [email protected] 31. Yves ROISIN Laboratoire de Biologie Animale et Cellulaire Service d'Eco-Ethologie Evolutive Université Libre de Bruxelles C.P. 160/12 50 av. F.D. Roosevelt 1050 Bruxelles Belgium Tel: +32 2 650 45 12 Fax: +32 2 650 24 45 Email: [email protected] 32. Tomas ROSLIN Dept. of Biological and Environmental Sciences University of Helsinki PO Box 65 00014 Helsinki Finland Tel: +358 9 191 57750 Fax: +358 9 191 57694 Email: [email protected]

33. Mirna SAMANIEGO Smithsonian Tropical Research Institute Apartado 0843-03092 Balboa Ancon Panama City Panama Tel: +507 212 8233 Fax: +507 212 8148 Email: [email protected] 34. Jürgen SCHMIDL Department of Ecology & Nature Conservation Institute for Zoology I University of Erlangen Staudtstr. 5 91058 Erlangen Germany Tel: +49 9131 8528076 Fax: +49 911 5188572 Email: [email protected] http://www.uni-erlangen.de/ 35. Line SORENSEN Zoological Museum University of Copenhagen Universitetsparken 15 2100 Copenhagen Ø Denmark Tel: +45 3532 1001 Fax: +45 3532 1010 Email: [email protected] 36. Neil SPRINGATE Department of Entomology The Natural History Museum Cromwell Road London SW7 5BD United Kingdom Tel: +44 207-942-5424 Email: [email protected] 37. Christian VAN OSSELAER Florinvest S.A., Biotrac SPRL 84 rue Stevens Delannoy 1020 Brussels Belgium Tel: +32.2.353.00.28 Fax: +32.2.353.05.81 Email: [email protected]








http://www.uni-erlangen.de/






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7. Statistical information on participants Disciplines represented Entomology, ecology; taxonomy; management (of international tropical research programme, NGO or private companies) ; statistics/modeling; botany. Distribution by nationality

Country Number 1 Australia 1 2 Belgium 5 3 Brasil 3 4 Czech Republic 1 5 Denmark 1 6 Finland 1 7 France 5 8 Germany 3 9 Italy 1 10 Luxembourg 1 11 New Zealand 2 12 Norway 1 13 Panama 4 14 Papua New Guinea 1 15 Switzerland 1 16 United Kingdom 6

37 Distribution by gender 6 females, 31 males Distribution by age groups

Age Number 20-30 3 30-40 10 40-50 22 50-60 2

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Participants in front of Manneken Pis, near Brussels' Grand Place, 8 July 2005

Documents

United Nations Environment Programme … · United Nations Environment Programme WORKSHOP The Last Biotic Frontier: Towards a Census of Canopy Life Scientific Report Royal Belgian