Workshops

Survival of Species in Fragmented Landscapes

Project Workshops

Summary of Ecological Modelling Workshop: 29th-31st January 1999 (Tvärminne)
At the end of January 1999, two teams of the FRAGLAND-TMR-Network, the Helsinki team (Ilkka Hanski, Rosemary Setchfield, Bob O’Hara, Mikko Heino, Martin Rost, Katrin Schöps and Weidong Gu) and the Leipzig team (Christian Wissel, Martin Drechsler, Karin Frank, Karin Johst and Thomas Stephan), came together in Tvärminne (Finland) to perform a common workshop on ecological modelling. The aim of this workshop was (1) to inform each other about the modelling approaches and techniques used in the individual groups to provide a common basis and (2) to look for opportunities of future cooperations on FRAGLAND- relevant problems. Each participant gave a presentation where great importance was laid to give enough time to come to a real understanding. The talks covered a large range of aspects relevant in the context of "Species survival and evolution in fragmented landscapes" and can be assigned to three main themes: Principles and methods of stochastic modelling; Parameter estimation; Dynamic landscapes. For a list of the talks see the table below. The most important results of the workshop among others are the following agreements concerning future collaborations between memebers from both teams: 1. Bob O’Hara will go to Leipzig to give a course on Bayesian statistics and the Monte Carlo simulation-based method of parameter estimation. This is important to get a common basis concering this component of model analysis. 2. Katrin Schöps (HKI) and Karin Johst (LPZ) will start a common project on a plant-insect system where a lot of field data are already available. 3. The metapopulation models used in Helsinki and Leipzig are found to have nearly the same funtional structure. It has been figured out that the existing methods for parameter estimation (HKI), prediction of metapopulation survival (LPZ) and decision analysis for metapopulation management (LPZ) fit very well together. A combination of these three components provides a modelling framework that allows empirical data to be analyzed in a way that a given landscape can be evaluated from the perspective of metapopulation survival and effective conservational strategies can be derived. Such a framework can be applied to any metapopulation system considered in the TMR-Network. Comparisons become possible with the result that the role of landscape fragmentation can be assessed. In a first step, Bob O’Hara, Atte Moilanen, Ilkka Hanski (all HKI), Karin Frank, Martin Drechsler and Christian Wissel (all LPZ) will try to apply this framework on the American pika metapopulation to test and demonstrate its potential. Details of the further collaboration will be discussed after the ESF-Meeting on Metapopulation Dynamics held in April 1999 in Tväminne/Finland.

Summary of Ecological Modelling Workshop: 29th-31st January 1999 (Tvärminne)

At the end of January 1999, two teams of the FRAGLAND-TMR-Network, the Helsinki team (Ilkka Hanski, Rosemary Setchfield, Bob O’Hara, Mikko Heino, Martin Rost, Katrin Schöps and Weidong Gu) and the Leipzig team (Christian Wissel, Martin Drechsler, Karin Frank, Karin Johst and Thomas Stephan), came together in Tvärminne (Finland) to perform a common workshop on ecological modelling.

The aim of this workshop was (1) to inform each other about the modelling approaches and techniques used in the individual groups to provide a common basis and (2) to look for opportunities of future cooperations on FRAGLAND- relevant problems. Each participant gave a presentation where great importance was laid to give enough time to come to a real understanding. The talks covered a large range of aspects relevant in the context of "Species survival and evolution in fragmented landscapes" and can be assigned to three main themes: Principles and methods of stochastic modelling; Parameter estimation; Dynamic landscapes. For a list of the talks see the table below.

The most important results of the workshop among others are the following agreements concerning future collaborations between memebers from both teams:

1. Bob O’Hara will go to Leipzig to give a course on Bayesian statistics and the Monte Carlo simulation-based method of parameter estimation. This is important to get a common basis concering this component of model analysis.
2. Katrin Schöps (HKI) and Karin Johst (LPZ) will start a common project on a plant-insect system where a lot of field data are already available.
3. The metapopulation models used in Helsinki and Leipzig are found to have nearly the same funtional structure. It has been figured out that the existing methods for parameter estimation (HKI), prediction of metapopulation survival (LPZ) and decision analysis for metapopulation management (LPZ) fit very well together. A combination of these three components provides a modelling framework that allows empirical data to be analyzed in a way that a given landscape can be evaluated from the perspective of metapopulation survival and effective conservational strategies can be derived. Such a framework can be applied to any metapopulation system considered in the TMR-Network. Comparisons become possible with the result that the role of landscape fragmentation can be assessed. In a first step, Bob O’Hara, Atte Moilanen, Ilkka Hanski (all HKI), Karin Frank, Martin Drechsler and Christian Wissel (all LPZ) will try to apply this framework on the American pika metapopulation to test and demonstrate its potential. Details of the further collaboration will be discussed after the ESF-Meeting on Metapopulation Dynamics held in April 1999 in Tväminne/Finland.

Christian Wissel	Standard methods for analyzing stochastic population models
Martin Drechsler	The range of applicability of presence-absence models
Karin Johst	How to handle temporally correlated stochasticity?
Karin Frank	Model simplification by using aggregation techniques
Bob O’Hara	Bayesian and Monte-Carlo methods of parameter estimation
Martin Drechsler	Conservational decision analysis in front of uncertainty
Mikko Heino	Evolution of dispersal
Katrin Schöps	Linking fieldwork and modelling (a new project)
Thomas Stephan	EXI: software for risk assessment of local populations
Ilkka Hanski	A new approach of modelling dynamic landscapes
Weidong Gu	Metapopulations in dynamic landscapes
Martin Rost	Pattern formation in a moth-parasitoid system
Rosemary Setchfield	Modelling of host-parasitoid dynamics

From my own view, I would say that the workshop was extremely stimulating. Each of us went home with a lot of new ideas. The chance of intensively thinking about the ideas of the partners opens up the opportunity of synergistic effects where everybody benefits from each other. This was a great experience and gave a lot of energy. By the way, the workshop took place in perfect surroundings (-30°C, snow, blue sky, sunshine, sauna, table tennis in the night, each day salmon - hm) so that everybody felt well - a mechanism to establish the network also from a social point of view.

Karin Frank, Leipzig

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Survival of Species in Fragmented Landscapes

TMR Workshops

Summary of Dispersal and Modelling Workshop: 21st February 1999 (Cordoba)
TMR FRAGLAND group, Córdoba 21 February 1999 Dispersal Workshop Chairperson : Chris D. Thomas Abstract prepared by Gabriel Nève (Córdoba) and Larissa Conradt (Leeds) Estimating dispersal rates One of the aims of the network is to get some meaningful comparisons of estimates of dispersal range between sites. This raises the question of standard fieldwork methods, and possible adjustments/corrections being applied to the resulting data, depending on the landscapes features, especially when behaviour is compared between large continuous habitats and fragmented habitats. ? Chris Thomas mentioned the following points : 1. Mark-Release-Recapture (MRR) results should be recorded on a grid square basis and with a standard catching protocol 2. Probability of recording different distances depends on the size of the study area (and on recapturing effort), and corrections are needed accordingly. 3. It is a great advantage for estimating dispersal rates and distances to use more than one method (e.g., release experiments and tracking of individuals). Again, there is need for standardisation of the methods. 4. One can never control for all variables (e.g., temperature, habitat quality, patch size and connectivity etc.), and comparisons between studies have to take this into account. ? With respect to release experiments, Paul Brakefield pointed out that wild caught individuals may display a different behaviour than laboratory-bred ones. This is a problem when estimating quantitative values of dispersal, but is less important if only relative differences between populations are investigated. Paul recommended to always release a cohort of wild caught butterflies as a control. ? MRR studies have limitations in estimating colonisation rates: in 5 releases into new habitat in the UK, colonisation rates were 10 times higher than expected from MRR studies. ? Peter Turchin suggested that crude dispersal distances from MRR data are not sufficient to estimate potential colonisation rates, because of the following confounding factors: - uneven sampling area - long distance attraction - landscape shape - temporal sampling period - temperature Therefore studies should be complemented by observations of individual behaviour, (in particular at habitat boundaries), and individual-based random walk models. ? Ilkka Hanski suggested that MRR studies should incorporate as many variables of the metapopulation approach as possible. In particular, attention should be given to the following parameters: - patch size - daily survival rate - daily emigration rate - connectivity - survival during migration - probability of migration. In specific studies, patch linked estimates should be given, and these should take into account local densities. ? Problems are raised about using metapopulation approaches for estimating dispersal and colonisation parameters in more continuous landscapes, and Peter Turchin emphasises the need for individual-based models. ? The need for, as well as the difficulty of, following individual butterflies to monitor their dispersal behaviour was discussed. ? It was suggested to use genetic markers to measure gene flow. Helsinki is trying to do this, but so far it has proved difficult to obtain sufficient polymorphic markers. Evolution of dispersal rates ? Selective pressures in different habitats can influence the evolution of dispersal rates. Affected traits could be: morphology, flight behaviour, speed, physiology, and behavioural responses to habitat. . ? Isabelle Olivieri summarised modelling approaches on the topic of evolution of dispersal rates: 1. Metapopulation models - stochastic models - deterministic models 2. Individual-based models 3. ESS models 4. Models including phenotypic plasticity It is necessary to investigate how much predictions differ between these models. Three selective pressures are likely to operate at the same time: - local adaptations (often favouring low dispersal rates) - inbreeding avoidance (favouring higher dispersal rates) - colonisation advantages / migration risks ? To investigate the evolution of dispersal rates in relation to habitat fragmentation, field experiments will be conducted, which compare the dispersal rates of individuals from the same species but different populations (stemming from habitats with different degrees of fragmentation). Another promising approach was the concurrent studies of species with the same host plant and similar habitat requirements in a given area (see Rob Wilson's talk). ? Paul Brakefield suggested selection experiments in greenhouses to investigate the evolution of dispersal abilities in Bicyclus, and wind tunnel experiments to correlate flight abilities to morphological traits. ? Particular attention should be given to traits related to dispersal behaviour in time (dormancy) or space, such as development time and some morphological traits. Hypothesis of behaviour based on spatial distribution of habitat patches should be tested against hypothesis of conspecific attraction (see Larissa Conradt’s talk, or "Measuring the degree of sexual segregation in group-living animals", J. Anim. Ecol. 67 : 217-226, 1998, and "Social segregation in red deer and feral soay sheep is not a by-product of habitat segregation", Animal Behaviour, in press, 1999). Modelling/Data collection Workshop Chairperson : Karin Frank Modelling requirements ? Karin Frank stressed that one has to first specify the biological question that one want to ask before one can decide what model and what data are needed. Karin summarised the steps of building a model, as follows: 1. What is the question? - Does one want to understand processes (mainly qualitative questions) or to make predictions (mainly quantitative questions)? - In the context of FRAGLAND, we will want to answer questions relating to evolution (understanding processes) and to conservation (making predictions, but also understanding processes). 2. From the perspective of the question: - Which are the important processes, factors, and interactions? - Which is the appropriate target quantity? - Which level of data resolution (patch occupancy, individual behaviour,...?) is needed/available? 3. Development of the model structures - starting from a relatively simple model structure that concentrates on most essential factors, details are included step by step (‘hierarchical approach’); generally, models should be as simple as possible but as complex as needed. - during the development of the model, it will become apparent which factors are crucial (therefore an accurate measurement is needed), and which factors are marginal (therefore it is sufficient to know their order of magnitude); this will influence the data collection process - at the same time the observations of processes by field ecologists should be taken into account to modify the model structure => modelling has to be a continuous process between modeller and field ecologist! Information that can be used for modelling: 1. Quantitative information - ‘hard data’ (direct measurements) - ‘weak data’ (direct estimates) - indirect estimates 2. Qualitative information - system knowledge (mechanisms, phenomena, patterns) 3. Possible scenarios - set of possible situations 4. Hypotheses 5. Patterns If data are missing or can only be approximately estimated, sensitivity analyses should be carried out: - to assess which factors are essential or marginal - to investigate worst/best case parameter values (‘reality’ is in between) - to compare extreme scenarios (e.g., clumped/evenly distributed landscape) Testing of hypotheses: Hypothesis => simulation => comparison with empirical patterns ?_________________________<________________________? feedback Comparability ? Is it possible to develop a general ‘standard model’ with a common level of resolution which can be geared to specific situations by sub-programs with details to estimate parameters? - points for discussion: What problems do we want to solve with such a model? How can we reach comparability? ? To decide about comparative data sampling/modelling, detailed discussions in small groups are necessary, and subsequent interactions between groups to preserve comparability. ? Spatial modelling approaches can be based 1. on a metapopulation structure (i.e., distinct patches in a matrix of unsuitable habitat) 2. on a lattice of grid cells 3. on a continuous landscape It is important to find modelling approaches which bridge the gap between these different modelling approaches. This problem should be addressed in parallel by modellers and field ecologists. ? Karin suggests as a first step towards bridging the gap between grid-based and metapopulation models to handle a metapopulation situation with grid-based methods and try to reconcile the approaches. ? Example of the problem of comparability: estimates of dispersal rates for Plebejus argus seem to differ between Holland, Spain and Britain by orders of magnitude, but a proper comparison is not currently possible with the available data. A special workshop was scheduled for the afternoon to discuss the problems of comparative estimation of dispersal rates. ? Ilkka Hanski reminded us that the software of the VM dispersal model is freely available on the internet at http://www.helsinki.fi/science/metapop/. ? Discussion about the right scale of grid solution.

Summary of Dispersal and Modelling Workshop: 21st February 1999 (Cordoba)

TMR FRAGLAND group, Córdoba
21 February 1999
Dispersal Workshop
Chairperson : Chris D. Thomas

Abstract prepared by Gabriel Nève (Córdoba) and Larissa Conradt (Leeds)

Estimating dispersal rates

One of the aims of the network is to get some meaningful comparisons of estimates of dispersal range between sites. This raises the question of standard fieldwork methods, and possible adjustments/corrections being applied to the resulting data, depending on the landscapes features, especially when behaviour is compared between large continuous habitats and fragmented habitats.

? Chris Thomas mentioned the following points :
1. Mark-Release-Recapture (MRR) results should be recorded on a grid square basis and with a standard catching protocol
2. Probability of recording different distances depends on the size of the study area (and on recapturing effort), and corrections are needed accordingly.
3. It is a great advantage for estimating dispersal rates and distances to use more than one method (e.g., release experiments and tracking of individuals). Again, there is need for standardisation of the methods.
4. One can never control for all variables (e.g., temperature, habitat quality, patch size and connectivity etc.), and comparisons between studies have to take this into account.

? With respect to release experiments, Paul Brakefield pointed out that wild caught individuals may display a different behaviour than laboratory-bred ones. This is a problem when estimating quantitative values of dispersal, but is less important if only relative differences between populations are investigated. Paul recommended to always release a cohort of wild caught butterflies as a control.

? MRR studies have limitations in estimating colonisation rates: in 5 releases into new habitat in the UK, colonisation rates were 10 times higher than expected from MRR studies.

? Peter Turchin suggested that crude dispersal distances from MRR data are not sufficient to estimate potential colonisation rates, because of the following confounding factors:
- uneven sampling area
- long distance attraction
- landscape shape
- temporal sampling period
- temperature
Therefore studies should be complemented by observations of individual behaviour, (in particular at habitat boundaries), and individual-based random walk models.

? Ilkka Hanski suggested that MRR studies should incorporate as many variables of the metapopulation approach as possible. In particular, attention should be given to the following parameters:
- patch size
- daily survival rate
- daily emigration rate
- connectivity
- survival during migration
- probability of migration.
In specific studies, patch linked estimates should be given, and these should take into account local densities.

? Problems are raised about using metapopulation approaches for estimating dispersal and colonisation parameters in more continuous landscapes, and Peter Turchin emphasises the need for individual-based models.

? The need for, as well as the difficulty of, following individual butterflies to monitor their dispersal behaviour was discussed.

? It was suggested to use genetic markers to measure gene flow. Helsinki is trying to do this, but so far it has proved difficult to obtain sufficient polymorphic markers.

Evolution of dispersal rates

? Selective pressures in different habitats can influence the evolution of dispersal rates. Affected traits could be: morphology, flight behaviour, speed, physiology, and behavioural responses to habitat. .

? Isabelle Olivieri summarised modelling approaches on the topic of evolution of dispersal rates:
1. Metapopulation models
- stochastic models
- deterministic models
2. Individual-based models
3. ESS models
4. Models including phenotypic plasticity
It is necessary to investigate how much predictions differ between these models.
Three selective pressures are likely to operate at the same time:
- local adaptations (often favouring low dispersal rates)
- inbreeding avoidance (favouring higher dispersal rates)
- colonisation advantages / migration risks

? To investigate the evolution of dispersal rates in relation to habitat fragmentation, field experiments will be conducted, which compare the dispersal rates of individuals from the same species but different populations (stemming from habitats with different degrees of fragmentation). Another promising approach was the concurrent studies of species with the same host plant and similar habitat requirements in a given area (see Rob Wilson's talk).

? Paul Brakefield suggested selection experiments in greenhouses to investigate the evolution of dispersal abilities in Bicyclus, and wind tunnel experiments to correlate flight abilities to morphological traits.

? Particular attention should be given to traits related to dispersal behaviour in time (dormancy) or space, such as development time and some morphological traits. Hypothesis of behaviour based on spatial distribution of habitat patches should be tested against hypothesis of conspecific attraction (see Larissa Conradt’s talk, or "Measuring the degree of sexual segregation in group-living animals", J. Anim. Ecol. 67 : 217-226, 1998, and "Social segregation in red deer and feral soay sheep is not a by-product of habitat segregation", Animal Behaviour, in press, 1999).

Modelling/Data collection Workshop
Chairperson : Karin Frank

Modelling requirements

? Karin Frank stressed that one has to first specify the biological question that one want to ask before one can decide what model and what data are needed. Karin summarised the steps of building a model, as follows:
1. What is the question?
- Does one want to understand processes (mainly qualitative questions) or to make predictions (mainly quantitative questions)?
- In the context of FRAGLAND, we will want to answer questions relating to evolution (understanding processes) and to conservation (making predictions, but also understanding processes).
2. From the perspective of the question:
- Which are the important processes, factors, and interactions?
- Which is the appropriate target quantity?
- Which level of data resolution (patch occupancy, individual behaviour,...?) is needed/available?
3. Development of the model structures
- starting from a relatively simple model structure that concentrates on most essential factors, details are included step by step (‘hierarchical approach’); generally, models should be as simple as possible but as complex as needed.
- during the development of the model, it will become apparent which factors are crucial (therefore an accurate measurement is needed), and which factors are marginal (therefore it is sufficient to know their order of magnitude); this will influence the data collection process
- at the same time the observations of processes by field ecologists should be taken into account to modify the model structure
=> modelling has to be a continuous process between modeller and field ecologist!
Information that can be used for modelling:
1. Quantitative information
- ‘hard data’ (direct measurements)
- ‘weak data’ (direct estimates)
- indirect estimates
2. Qualitative information
- system knowledge (mechanisms, phenomena, patterns)
3. Possible scenarios
- set of possible situations
4. Hypotheses
5. Patterns
If data are missing or can only be approximately estimated, sensitivity analyses
should be carried out:
- to assess which factors are essential or marginal
- to investigate worst/best case parameter values (‘reality’ is in between)
- to compare extreme scenarios (e.g., clumped/evenly distributed landscape)

Testing of hypotheses:

Hypothesis => simulation => comparison with empirical patterns
?_________________________<________________________?
feedback

Comparability

? Is it possible to develop a general ‘standard model’ with a common level of resolution which can be geared to specific situations by sub-programs with details to estimate parameters?
- points for discussion: What problems do we want to solve with
such a model?
How can we reach comparability?

? To decide about comparative data sampling/modelling, detailed discussions in small groups are necessary, and subsequent interactions between groups to preserve comparability.

? Spatial modelling approaches can be based
1. on a metapopulation structure (i.e., distinct patches in a matrix of unsuitable habitat)
2. on a lattice of grid cells
3. on a continuous landscape
It is important to find modelling approaches which bridge the gap between these different modelling approaches. This problem should be addressed in parallel by modellers and field ecologists.

? Karin suggests as a first step towards bridging the gap between grid-based and metapopulation models to handle a metapopulation situation with grid-based methods and try to reconcile the approaches.

? Example of the problem of comparability:
estimates of dispersal rates for Plebejus argus seem to differ between Holland, Spain and Britain by orders of magnitude, but a proper comparison is not currently possible with the available data.
A special workshop was scheduled for the afternoon to discuss the problems of comparative estimation of dispersal rates.

? Ilkka Hanski reminded us that the software of the VM dispersal model is freely available on the internet at http://www.helsinki.fi/science/metapop/.

? Discussion about the right scale of grid solution.

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Survival of Species in Fragmented Landscapes

TMR Workshops

Summary of Effective Population Size Workshop: 21st February 1999 (Cordoba)
1. Accurate knowledge of the effective population sizes of the populations/metapopulations is a critical parameter for extending ecological studies to genetical models, yet there are very few direct estimates of the relationship between census and effective population size in natural populations. 2. There is potential for using molecular markers (microsatellites) in combination with MRR studies to obtain much needed direct estimates of Ne. 3. However, in highly subdivided metapopulations, such as the Aland cinxia, where inbreeding is high (and local genetic variability low), large numbers of marker loci would be needed to distinguish parental genotypes. 4. It may be useful to expand Bicyclus cage experiments to measure Ne (variances in reproductive success, incl. sperm competiton), for example at different densities, though the relevance of these estimates to any natural population is questionable. Of the field systems studied within Fragland, the Melitaea cinxia metapopulation in Aland is possibly the most suitable for a study of effective population size. However, Ilkka Hanski thought that while such a study was feasible, albeit with a substantial workforce, it would be wiser to first develop more microsatellite markers. Jeremy Thomas suggested that, in the absence of sufficient detectable molecular marker variation, another option to getting a rough estimate of the variance in reproductive success would be to follow individual female butterflies, and base estimates of their reproductive success on the suitability of microhabitat where they lay eggs. M. cinxia, H. comma and P. argus are impractical for such a study, but it might be achievable with Maculinea nausithous or Cupido minimus.

Summary of Effective Population Size Workshop: 21st February 1999 (Cordoba)

1. Accurate knowledge of the effective population sizes of the populations/metapopulations is a critical parameter for extending ecological studies to genetical models, yet there are very few direct estimates of the relationship between census and effective population size in natural populations.

2. There is potential for using molecular markers (microsatellites) in combination with MRR studies to obtain much needed direct estimates of Ne.

3. However, in highly subdivided metapopulations, such as the Aland cinxia, where inbreeding is high (and local genetic variability low), large numbers of marker loci would be needed to distinguish parental genotypes.

4. It may be useful to expand Bicyclus cage experiments to measure Ne (variances in reproductive success, incl. sperm competiton), for example at different densities, though the relevance of these estimates to any natural population is questionable.

Of the field systems studied within Fragland, the Melitaea cinxia metapopulation in Aland is possibly the most suitable for a study of effective population size. However, Ilkka Hanski thought that while such a study was feasible, albeit with a substantial workforce, it would be wiser to first develop more microsatellite markers.

Jeremy Thomas suggested that, in the absence of sufficient detectable molecular marker variation, another option to getting a rough estimate of the variance in reproductive success would be to follow individual female butterflies, and base estimates of their reproductive success on the suitability of microhabitat where they lay eggs. M. cinxia, H. comma and P. argus are impractical for such a study, but it might be achievable with Maculinea nausithous or Cupido minimus.

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Survival of Species in Fragmented Landscapes

TMR Workshops

Melitaea cinxia workshop: 21st February 1999 (Cordoba)
The aim of the workshop was to preliminarily plan an experiment on detecting differences in migration rates among metapopulations of M. cinxia with different degrees of fragmentation. We predict that the selective pressures for optimal migration rates differ according to the degree of habitat fragmentation. This would lead to differences in the evolution of migration rate. The experiment will include individuals from three habitat structures: (1) Relatively non-fragmented or continuous habitat, (2) fragmented habitat with large and isolated habitat patches, and (3) fragmented habitat with small and non-isolated habitat patches. Individuals for the experiment will be collected as larvae (probably 500 larvae from each habitat type), reared to adult stage and released, individually marked, within a network of unoccupied but suitable habitat patches covering about 4 km2 in eastern Åland, SW Finland. The suitability of the patch network can be checked prior to the actual experiment with the virtual metapopulation model. Before releasing, individuals will be photographed (for fluctuating asymmetry of wings, and for wing pattern, size and shape), weighed (for calculation of wing loading), and a small piece of wing will be sampled for DNA-analyses. During the experiment, all patches in the area will be visited daily with high and constant effort to recapture butterflies. These data reveal for example emigration and immigration rates, migration distances, stay-times, adult longevity, and effects of population density and patch shape on migration rate. Moreover, some researchers concentrate on the behaviour of butterflies. There are two aims in the behavioural study: (1) To observe the behaviour close to and at the patch boundary which is very important to the emigration rate, and (2) to observe the behaviour during migration, i.e. between habitat patches. For example, the turning angles and frequencies will probably differ from each other and from what occur within habitat patches. Butterflies hardly leave the patches as often as they encounter the patch boundary, but will instead frequently turn back to the patch. On the other hand, when flying on a matrix habitat butterflies probably use more directed flight than within patch. Later in the autumn, all larval groups will be collected from the field for DNA-analyses. These analyses should enable the detection of parents of each larval group, which gives further information on the movements of butterflies within the patch network, as well as on spatial allocation of the offspring and breeding success of individuals originating from different habitat structures.

Melitaea cinxia workshop: 21st February 1999 (Cordoba)

The aim of the workshop was to preliminarily plan an experiment on detecting differences in migration rates among metapopulations of M. cinxia with different degrees of fragmentation. We predict that the selective pressures for optimal migration rates differ according to the degree of habitat fragmentation. This would lead to differences in the evolution of migration rate.

The experiment will include individuals from three habitat structures: (1) Relatively non-fragmented or continuous habitat, (2) fragmented habitat with large and isolated habitat patches, and (3) fragmented habitat with small and non-isolated habitat patches. Individuals for the experiment will be collected as larvae (probably 500 larvae from each habitat type), reared to adult stage and released, individually marked, within a network of unoccupied but suitable habitat patches covering about 4 km2 in eastern Åland, SW Finland. The suitability of the patch network can be checked prior to the actual experiment with the virtual metapopulation model.

Before releasing, individuals will be photographed (for fluctuating asymmetry of wings, and for wing pattern, size and shape), weighed (for calculation of wing loading), and a small piece of wing will be sampled for DNA-analyses. During the experiment, all patches in the area will be visited daily with high and constant effort to recapture butterflies. These data reveal for example emigration and immigration rates, migration distances, stay-times, adult longevity, and effects of population density and patch shape on migration rate. Moreover, some researchers concentrate on the behaviour of butterflies. There are two aims in the behavioural study: (1) To observe the behaviour close to and at the patch boundary which is very important to the emigration rate, and (2) to observe the behaviour during migration, i.e. between habitat patches. For example, the turning angles and frequencies will probably differ from each other and from what occur within habitat patches. Butterflies hardly leave the patches as often as they encounter the patch boundary, but will instead frequently turn back to the patch. On the other hand, when flying on a matrix habitat butterflies probably use more directed flight than within patch.

Later in the autumn, all larval groups will be collected from the field for DNA-analyses. These analyses should enable the detection of parents of each larval group, which gives further information on the movements of butterflies within the patch network, as well as on spatial allocation of the offspring and breeding success of individuals originating from different habitat structures.

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Survival of Species in Fragmented Landscapes

TMR Workshops

"Explanation and demonstration of models of movement behaviour and evolution of migration rate" 2.-6.2. 2000 (Tvärminne)
Preliminary information
Participants
Draft program
How to find the Department, (Map2)

Preliminary information:
As we agreed in the Steering Committee meeting in Leiden, we will organize a 5-day workshop/meeting focused on modelling of movement behaviour and migration, including modelling the evolution of migration. There is much work going on in this area in Fragland, and it was considered that another opportunity to discuss these matters in detail would be very helpful. This would allow an opporunity for modellers to describe in detail the assumptions and structure of their models. We will discuss model predictions, and how they can be tested with existing data and how new experiments should be planned. Modelling of movement behaviour will include modelling at the individual level. A more detailed plan of exactly what we will cover in the workshop will be prepared in advance - all ideas most welcome! It would be good to know as early as possible who will participate.
Practical information:

Tvärminne Zoological Station

Accommodation costs: 280 FIM/night including all the meals (1999).

The workshop starts on Wednesday morning (2.2.), so it would be desired if you could arrive on Tuesday before 6 PM (the bus leaves at 6). Let me know the time of your arrival. There will also be a bus from Tvärminne to Helsinki on Sunday at 1 PM (driving time to Helsinki is approx. 2 hours).

There are only limited amount of rooms which means everybody can not have a single room. If you want to share a room, let me know.

The bus between the Airport and Helsinki takes half an hour.

Participants:
Rosemary Setchfield, Leipzig
David Gutierrez, Cordoba
Henri Leturque, Montpellier
Ido Pen, Montpellier
Tomas Roslin, Helsinki
Marko Nieminen, Helsinki
Ilkka Hanski, Helsinki
Bob O'Hara, Helsinki
Otso Ovaskainen, Helsinki
Mikko Heino, Helsinki
Niklas Wahlberg, Helsinki
Larissa Conradt, Leeds
Ted Bodsworth, Leeds
Adam Simmons, Leeds
Rob Wilson, Leeds
Dale Taneyhill, Leeds
Chris Thomas, Leeds
Jane Hill, Leeds
Nicolas Schtickzelle, Louvain
Gwennaelle Mennechez, Louvain
Eric Leboulenge, Louvain
Ilik Saccheri, Leiden

DRAFT PROGRAM

Concept: we have talks, presentations, demonstration and discussions all day from 9 am until 5 pm. After dinner, there is an opportunity to have sauna, smaller groups of people may wish to talk to each other, or we may decide to arrange some round table discussions.

TUESDAY 1 February
6 PM Bus to Tvärminne

WEDNESDAY 2 February

Morning

Introduction to workshop (Ilkka, 15 min)
Theme 1: Modelling of individual movement behavior

Introduction to the theme (Ilkka, 15 min)
Presentations by Rosemary and Larissa (you two could coordinate your presentations if that would be helpful; total time 2 hrs)

Afternoon
VM model

Introduction and model description (Ilkka, 1 hr)
Software demonstration (Niklas, 30 min)
Applications, Niklas and Michel (30 min each)

Genetic consequences of migration in small populations: a lab experiment (Ilik, 1 hr)

THURSDAY 3 February

Morning

Individual presentations from Helsinki (Tomas), Leeds, Louvain:
PLEASE SUPPLY NAMES OF SPEAKERS AND TITLES!!!

Afternoon
Theme 2: Evolution of migration rate
Introduction to the theme (Chris, 15 min)
General models, someone describing the work done in Montpellier (Who?,1 hr)
An individual-based model (Mikko, 1 hr)
Discussion: how do different models differ, what are their strengths and weaknesses, how could they be improved?

FRIDAY 4 February

Morning
Theme 2 continues
Empirical studies by Chris and others (2 hrs)
A field experiment (Ilkka, 1 hr)

Afternoon
Theme 3: Metapopulation modelling
Introduction to the theme (Ilkka, 15 min)
IFM, description and demonstration, by Bob and Atte (3 hrs)

SATURDAY 5 February

Morning

Field trip (bring warm clothing)

Afternoon
Theme 3 continues
Bayesian analysis of metapopulation data (Bob, 1 hr)
Meta-X, description and demonstration of the model, by Karin and Rosemary (2 hrs)

SUNDAY 6 February

Morning

Presentations on modelling metapopulation dynamics, Otso, Karin

Closing of the workshop

Lunch

1 PM, Departure to Helsinki and the airport

3 PM Bus in Helsinki

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Survival of Species in Fragmented Landscapes

TMR Workshops

"GIS and spatial modelling" 8.-13.1. 2001 (Louvain)

Participants
Practical info
Map of Louvain-La-Neuve
Program of Open Meeting
Guest speakers

SUMMARY

Preliminary info:
GIS COURSES: 8-9-10-11 January
This session will provide to participants with a basic (theoretical and practical) knowledge of GIS. Courses will be given by ESRI experienced trainers. The number of participants is unfortunately limited! So, TMR postdocs, task leaders and people directly involved in Fragland will have priority.
A small financial contribution (including lunches and informatics fees) is required : 125 EURO/participant.
OPEN MEETING: 12-13(-14?) January
This session will be made of courses, presentations and demonstrations on applications of GIS to population ecology. Different topics will be tackled as landscape analysis, habitat suitability analysis, spatial population dynamics modelling.

Participants of the GIS course (8-9-10-11 January):

Cordoba (2)
- Adrian Seymour
- David Guiterrez
Montpellier (3)
- Lucille Lafuma
- Sandrine Maurice
- Eric Imbert
Louvain-la-Neuve (12)
- Michel Baguette
- Gwénaëlle Mennechez
- Nicolas Schtickzelle
- Sabine Soetewey
- Sofie Vandewoestijne
- Virginie Stevens
- Eric le Boulangé
- Nicolas Titeux
- Marianne Schlesser
- Carine Petit
- Pierre Defourny
- Vincent Guissard

Leiden (1)
- Mathieu Joron
Leipzig (4)
- Rosemary Setchfield
- Stephanie Schadt
- Eloy Revilla
- Thorsten Wiegand
Helsinki (3)
- Mika Siljander
- Tarja Salmi
- Marko Nieminen
Leeds (5)
- Zoe Davies
- Ted Bosworth
- David Blakeley
- Deborah Sazer
- Stephen Hartley

Accommodation
Bed and Breakfast accommodation will be at the Relais in Louvain-la-Neuve. There is only one hotel in Louvain-la-Neuve, so we have to book hotel rooms for participants in GIS course (night:7-8-9-10-11) and for participants in the open meeting (day:12,13,14 morning).
Let us know rapidly who intend to participate in the open meeting. When will you leave Louvain-la-Neuve (14 or 15 January)?
Practical information: people who will not attend the GIS course but want to participate in the open meeting have to arrive on 11 January.
Excursion
We are planning an excursion on Sunday afternoon (14 January). Let us know whether you prefer a cultural excursion (Brugges? Bruxelles?) or bird watching on the seaside.
Thank you very much.
Best regards,
Michel, Gwen.
More information: Gwénaëlle Mennechez

Guest speakers
Jana Verboom
Department of Landscape Ecology, Alterra Green World Research, NL
Frank Adriaensen
Laboratory of Animal Ecology, University of Antwerp, Belgium
Ruth Swetnam
Ecological Systems Modelling Group, Centre for Ecology & Hydrology, Monks Wood, UK

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