Ph.D. position for a biologist at the Arctic Centre of the ... · The PhD candidate will be investigating dynamics and structure of an Arctic terrestrial food web, with geese as the

Ph.D. position for a biologist at the Arctic Centre

of the University of Groningen

This document contains: Letter of instruction

The job announcement

Project proposal

Referee report

Rebuttal

Letter of instruction

Dear Applicant,

This letter presents guidelines for your application for a research post leading to a PhD at the

Arctic Centre n the Graduate School for the Humanities. All posts are in general for a period

of four years.

The goal of the application format is to enable us to compare the relative merits of the

applicants efficiently and with a minimum of bias.

Please respect these guidelines. Applicants that do not follow this format might be rejected.

Your application should have the following elements:

1. A motivation letter.

2. A curriculum vitae (including date and place of birth, the name of your secondary

school, and your academic record, also -if applicable- titles of publications).

3. A certified copy of your college transcript (‘doctorandus’ or MA degree, or an

equivalent), as well as a list of examination results.

4. Email- and postal addresses of two academic referees.

5. A reflection on your expertise in:

- writing scientific papers

- analyzing large datasets

- using statistical (R, SPSS), survival (MARK) and/or spatial (GIS) software

- modelling and programming

- laboratory analysis and molecular techniques

- field work

6. Please do provide evidence of the level at which you master your skills

Do please send your entire application in only two pdf-files (one for the motivation letter,

and one for all the other documents) via the following web address:

http://www.rug.nl/about-us/work-with-us/job-opportunities/phd-positions

and do adhere to the application deadline in the advert. Late applications will not be taken

into consideration.

We plan to invite applicants for an interview within two weeks after the submission deadline.

We have no option to postpone the starting date after 1 May 2014.

PhD position for a biologist at the Arctic Centre (1,0 fte) (214063)

Organisation

Since its foundation in 1614, the University of Groningen has established

an international reputation as a dynamic and innovative university offering high-quality teaching and research. Its 27,000 students are encouraged to

develop their own individual talents through challenging study- and career paths. The University of Groningen is an international center of

knowledge: It belongs to the best research universities in Europe and is

allied with prestigious partner universities and networks worldwide.

Job description

The Arctic Centre is seeking a PhD candidate for the project ‘From historical data to a prediction of the future for geese on Arctic tundra’. The

project is part of the New Netherlands Polar Programme ‘Anthropogenic impact on Svalbard ecosystems’, funded via NWO, the Netherlands

Organisation for Scientific Research. The Arctic Centre has an inter-disciplinary research focus on long-term human-environment relations in

polar regions.

The PhD candidate will be investigating dynamics and structure of an

Arctic terrestrial food web, with geese as the central element. The Ph.D. project focuses on the analysis of historical data and a multi-year dataset

of plant-herbivore interaction and dynamics of predation into models of resource utilization, habitat selection and population dynamics. Field

experiments and molecular techniques are planned to support a better understanding of goose numbers and goose distribution on Arctic tundra

vegetation. For a full description of the project and the desired format of the application see: http://www.rug.nl/staff/m.j.j.e.loonen/projects

The Ph.D. candidate is expected to:

• Complete a Ph.D. dissertation based on four peer-reviewed articles (in English) within the four year period

• Participate in local research meetings and Ph.D. training • Participate in meetings with international experts and present

papers at international conferences

Qualifications

• Master degree in biology, preferably ecology

• High motivation to analyze field data into scientific papers • Proven skills in database use, statistical analysis and modelling

• Excellent command of English (oral and written) • Ability to work in a team

• Interested in doing interdisciplinary research

Conditions of employment

The University of Groningen offers the PhD a salary of € 2,083 (scale 10, number 0) gross per month in the first year, up to a maximum of € 2,664

(scale 10, number 3) gross per month in the final year, based on a full-time position. The position requires residence in Groningen, 38

hours/week research and research training, and must result in a PhD dissertation. The successful candidate will first be offered a temporary

position of 1,5 years with the perspective of prolongation for another 2,5 years. After 12 months there will be an evaluation on the perspectives of

the successful completion of the PhD thesis within the next 2,5 years. If these perspectives are poor, the contract may not be renewed.

The ultimate starting date is 1 May 2014.

Successful applications include a letter containing your motivation for

applying for this position (this includes a short description of how you

intend to set up the research) and a description of previous relevant research experience, a full curriculum vitae, a copy of diplomas and a list

of grades, the names and e-mail addresses of two academic referees and the vacancy number. Applications that are incomplete or are otherwise

faulty will not be taken into account.

Applications must be submitted electronically in PDF format by means of the application form, in two files: one for the letter and one for the cv and

other documents. The deadline for submission is 6 March 2014. Submission via

http://www.rug.nl/about-us/work-with-us/job-opportunities/phd-positions

The university is an equal opportunities employer. Because women are still underrepresented in a number of fields, they are particularly

encouraged to apply.

Information

For information please contact Dr. Maarten Loonen: [email protected]

New Netherlands Polar Programme Proposal form 2012

Page 1 of 11

The maximum length of a proposal is 11 pages

1. Details of proposal

Theme: Ο Ice, climate and sea level

Ο Polar oceans X Polar ecosystem Ο Humanities and changes in polar regions

Title: From historical data to a prediction of the future for geese on arctic tundra? Summary (max. 250 words):

This project studies the dynamics in the structure of an Arctic terrestrial food web and the genetic structure of a population of geese. Geese play a central role in this food web. Historical patterns of habitat distribution will be analysed to recognize the effect of local and global anthropogenic drivers of change. Observations over a time-series of 23 years on three trophic levels with details on food availability, diet selection, intake rate, plant-animal interaction, habitat use, reproductive success and timing of marked individuals (plants, geese and foxes) are combined with predator abundance, predation pressure and disturbance into a spatial model of resource utilisation, habitat distribution and population dynamics. Information on the genetic structure of the barnacle goose population will provide novel insight in contemporary patterns of divergence, population size and gene flow. While studying species-specific plant-animal interaction and immunological parameters we hope to understand historical changes in species distribution. All elements combined should enable the prediction of future scenarios of anthropogenic impact on species interaction within the food web.

2. Details of applicant

Name: Dr. Maarten J.J.E. Loonen Institution: University of Groningen, Arctic Centre Position: Ο Professor Ο Associate professor (UHD) X Assistant professor (UD)

Ο Other: Permanent position: X Yes Ο No, end date contract:

E-mail: [email protected] 2a. Alternative contact

Name: Prof. Dr. L. Hacquebord Tel: 050-363 6832 Email: [email protected]

3. Renewed application? Ο Yes X No In case of a renewed application please indicate the file number of the previous application and summarize the main changes

4. Applying for: X PhD student Ο Post Doc

5. Core Subsidy

5a. Please enlist peer-reviewed publications, international evaluations and scientific honours for polar research accomplished by you and your department/ research group

The current researchers of the Arctic Centre has (co)authored about 150 peer reviewed publications dedicated to polar topics. Within the Arctic Centre there are currently 9 PhD projects and 15 PhD theses have been finished. The Arctic Centre had a recent assessment over the period 2004-2009 according to the national Standard Evaluation Protocol 2009-2015 (SEP). Quote: The research quality at the Arctic Centre was described as unique and important, both for The Netherlands and internationally. The Polar Regions are virtual laboratories for global change as well


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as adaptation and human impacts on the environment. This circumstance calls for high-quality interdisciplinary research and Arctic Centre is well positioned to continue this commitment. The Arctic Centre has maintained a high level of quality in its research across a broad range of fields. Quote: The work of the Arctic Centre makes an internationally important contribution to heritage and wildlife management of fragile Arctic ecosystems including such internationally important topics as bird populations, fish stocks and whaling. The work of the biological archaeologists contributes to an understanding of the past history and ecology of plant and animal species and this is of relevance in nature conservation. Quote: The two PRC members who were not previously so aware of it, were surprised and impressed by the exciting interdisciplinary work of the Arctic Centre and the way it combined archaeology with other fields. Quote: The vitality of the Arctic Centre appears excellent. The list of literature cited in this proposal contains 13 peer-reviewed papers by Loonen. Some additional peer-reviewed papers regarding polar research: Dolnik, OV, Metzger BJ & Loonen MJJE (2011) Keeping the clock set under the midnight sun: diurnal

periodicity and synchrony of avian Isospora parasites cycle in the High Arctic. Parasitology 138 (9): 1077-1081.

Hahn S, Loonen MJJE & Klaassen M (2011) The reliance on distant resources for egg formation in high Arctic breeding barnacle geese Branta leucopsis. J. Avian Biol. 42: 159-168.

Prestrud KW, Åsbakk K, Fuglei E, Mørk T, Stien A, Ropstad E, Tryland M, Gabrielsen GW, Lydersen, C, Kovacs KM, Loonen MJJE, Sagerup K, & Oksanen A (2007) Serosurvey for Toxoplasma gondii in arctic foxes and possible sources of infection in the high Arctic of Svalbard. Vet. Parasitol. 150: 6-12.

Dolnik OV & Loonen MJJE (2007) Isospora plectrophenaxia n. sp (Apicomplexa: Eimeriidae), a new coccidian parasite found in Snow Bunting (Plectrophenax nivalis) nestlings on Spitsbergen. Parasitology Research 101: 1617-1619.

Pezzanite B, Rockwell RF, Davies JC & Loonen MJJE & Seguin RJ (2005) Has habitat degradation affected foraging behaviour and reproductive success of lesser snow geese (Chen caerulescens caerulescens)? Ecoscience 12: 439-446.

Stahl J, Tolsma PH, Loonen MJJE & Drent RH (2000) Subordinates explore but dominants profit: resource competition in high arctic barnacle goose flocks. Animal behaviour 61: 257-264.

Loonen MJJE, Bruinzeel LW, Black JM & Drent RH (1999) The benefit of large broods in Barnacle geese: a study using natural and experimental manipulations. J. Anim. Ecol. 68: 753-768.

Bakker C & Loonen MJJE (1998) The influence of goose grazing on the growth of Poa arctica: overestimation of overcompensation. Oikos 82: 459-466.

Wal R van der & Loonen MJJE (1998) Goose droppings as food for reindeer. Can. J. Zool. 76: 1117-1122.

Loonen MJJE, Oosterbeek K, Drent RH (1997) Variation in growth of young and adult size in barnacle geese Branta leucopsis: evidence for density dependence. Ardea 85: 177-192.

Bishop CM, Butler PJ, El Haj AJ, Egginton S & Loonen MJJE (1996) The morphological development of the locomotor and cardiac muscles of the migratory barnacle goose (Branta leucopsis). J. Zool., Lond. 239: 1-15.

Williams TD, Loonen MJJE & Cooke F (1994) Fitness consequences of parental behavior in relation to offspring number in a precocial species: the lesser snow goose. The Auk 111(3): 563-572.

5b. Please enlist long- term investment to infrastructure as a sign of commitment to polar research from your research institution

Since 1970, the Arctic Centre of the University of Groningen is committed to polar research. It is a national platform for polar research. It has been networking (starting the Willem Barentsz Polar Institute as a national organization for polar researchers) and creating initiatives for the Dutch polar science community by organizing scientific expeditions to Spitsbergen (Smeerenburg, Nordenskiøldkysten, Edgeøya). It has a leading role in polar public outreach (Poolnacht, national symposia, press information, arcticstation.nl). The Arctic Centre is running the Netherlands Arctic Station in Ny-Ålesund and has developed and maintained a local ecological research program for 23 years. The Arctic Centre is committed to an international representation in NySMAC, working groups AMAP and CAFF of the Arctic Council and IASC. We made the University of Groningen partner in SIOS, an EU-ESFRI program for international


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scientific infrastructure with the aim to develop Svalbard to a regional Arctic monitoring site for system Earth. During the International Polar Year, the Arctic Centre has been coordinator of two international research projects (LASHIPA by Hacquebord, BIRDHEALTH by Loonen). Loonen has been coordinator of a workpackage in the EU-funded project FRAGILE, which studied plant-animal interaction on High Arctic tundra. Members of the Arctic Centre have been participating in all national polar committees. Our newest initiative is a 100 person multidisciplinary scientific expedition to Egeøya, planned in 2014 and financed by sponsors. At present this initiative is running in the finals for the national science outreach competition (Academische Jaarprijs). The University of Groningen is committed to the future of the Arctic Centre. Recently the Arctic Centre has received an increase in staff (4 fte and 3 Ph.D.) and replacement for the soon retiring Prof. Hacquebord is under selection. Annually 30 students follow courses at the Arctic Centre as part of their degree.

6.Composition of the research group List all staff members involved in the proposed research: provide name, initials, titles and type of involvement, e.g. daily guidance, technician, thesis supervisor, advisor. Name and title Specialization Institution Involvement hrs/wk

Dr MJJE Loonen arctic ecology University of Groningen, AC

thesis supervisor, daily guidance

6

PhD, TBA ecologist University of Groningen, AC

project researcher 40

Prof Dr P Palsbøl genetic markers population size

University of Groningen, CEES

advisor, laboratory facilities 2

Prof Dr J Komdeur

life history genomics

University of Groningen, CEES

advisor, laboratory facilities 2

Prof Dr AGJ Buma

polar phytoplankton

University of Groningen, ESRIG

laboratory facilities, training p.m.

Dr. J Prop goose ecology University of Groningen, AC

advisor 4

Dr. BA Nolet modeling herbivore behaviour

NIOO, Wageningen advisor p.m.

Prof Dr A Gröne pathology University Utrecht, Veterinary Medicine

biopsies, training p.m.

Dr. GW Gabrielsen

arctic ecotoxicology

Norwegian Polar Institute

advisor, link to NPI datasets and researchers

p.m.

Dr. A.J.M. Scheepstra

coordination / organization

Willem Barentsz Polar Institute

organizing workshops, overarching PhD and Postdoc projects

2

7. Summary for the general public (in Dutch, max. 100 words)

Veel ganzensoorten zijn door menselijk handelen sterk in aantal toegenomen. Grote aantallen overwinterende ganzen trekken in de zomer naar het noordpoolgebied en zouden daar dramatische effecten op de arctische toendra hebben. In de loop der jaren is veel kennis verzameld, maar het ontbreekt aan een coherent beeld om de langere termijn effecten van deze herbivore vogels te voorspellen. In dit project worden de unieke lange tijdsreeksen en monsters, die reeds verzameld zijn, geanalyseerd en aangevuld met doelgerichte experimenten om de verschillen tussen ganzensoorten te kwantificeren. Doel is om de effecten van ganzen op het terrestrische ecosysteem te beschrijven, begrijpen en voorspellen.

8. Top 5 publications of the research group related to the proposed research


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1. Kuijper DPJ, Ubels R & Loonen MJJE (2009) Density-dependent switches in diet: a likely mechanism for negative feedbacks on goose population increase? Polar Biology 32: 1789-1803.

2. Peery MZ, Beissinger SR, House RF, Bérubé M, Sellas A, and Palsbøll PJ (2008). Characterizing source-sink dynamics with genetic parentage assignments. Ecology 89:2746-2759

3. Richardson DS, Komdeur J, Burke T & von Schantz T (2005) MHC-based patterns of social and extra-pair mate choice in the Seychelles warbler. Proc. R. Soc. B 272: 759-767.

4. Kurvers RHJM, Prins HHT, van Wieren SE, van Oers K, Nolet BA, and Ydenberg RC (2010) The effect of personality on social foraging: shy barnacle geese scrounge more. Proceedings of the Royal Society, London B 277:601-608.

5. Letcher RJ, Bustnes JO, Dietz R, Jenssen BM, Jorgensen EH, Sonne C, Verreault J, Vijayan MM & Gabrielsen GW (2010) Exposure and effects assessment of persistent organic pollutants in arctic wildlife and fish. Science of the Total Environment 408: 2995-3043.

9. Description of the proposed research (4 pages maximum, including figures, excluding literature references, font size at least 10 points) Include details of objectives, innovative aspects, scientific approach, impact, necessity to work in the Arctic, and literature references

The proposed study makes use of existing data sets, novel analyses and continued field work to understand the dynamics in Arctic goose breeding success as an important component of the terrestrial Arctic food web. The project will provide building blocks for modeling future scenarios for goose demography on the Arctic tundra. The proposal is written as a multi-faceted contribution to the core program of the Arctic Centre. It will place the Ph.D. candidate in a central position linked to several of the other proposals in the core program and opens possibilities for close cooperation across a variety of research disciplines. Objectives are numbered 1 to 4. 1. Develop a spatial explicit distribution model for grazing pressure based on trophic interactions and anthropogenic drivers

Historical numbers of arctic goose populations are largely unknown, but an exponential increase in abundance since the middle of the last Century is well documented (Fox et al. 2010). The cause of this increase lies outside the Arctic and is largely related to human-induced changes on wintering grounds at lower latitudes (Jefferies et al. 2004). The consequences of the increase in goose numbers for the vegetation on the Arctic tundra range from substantial (Speed et al. 2009) to destructive (Kotanen & Jefferies 1997) and were studied in detail using captive geese in the FRAGILE project (O’Connell et al. 2006). Experimental manipulation of grazing pressure revealed effects of geese on carbon stocks in the soil (Sjögersten et al. 2010a), nutrient dynamics (Loonen & Solheim 1996, Zielke et al. 2004, Sjögersten et al. 2010b) and the competition between moss and grass (Van der Wal et al. 2001). Geese are an important food source to predators (Stahl & Loonen 1998, Ebbinge & Spaans 2002, Drent & Prop 2008) and impact lake ecosystems with their faeces (Van Geest et al. 2007). All in all, geese may be viewed as key species in the terrestrial Arctic ecosystem.

Within our study area, human activity affects goose distribution. The main goose brood rearing grounds are close to the village of Ny-Ålesund, where the vegetation is further enriched by nutrient additions and polluted by coal mining. Also the presence of predators like the polar bear, has been affected by humans through legislation and protective measures.

The barnacle goose (Branta leucopsis) population in Kongsfjorden started as a newly founded colony in 1980 and rapidly grew during periods of reduced predation pressure. As a result, the vegetation became heavily grazed (Loonen 1997, Loonen et al. 1998). However, since 2000, breeding success has been extremely low due to several consecutive years with high a predation pressure (mainly by arctic foxes), which in turn allowed the vegetation to recover. Over a period of 23 years, data on habitat use, timing of reproduction, breeding success, survival, grass growth, vegetation composition, predation pressure, predator populations and the population size of reindeer (the only other herbivore present) were collected. These data provide a unique opportunity for a detailed description of the consequences of anthropogenic drivers on the dynamics of a terrestrial arctic food web.

We will develop a spatial explicit distribution model (Baveco et al. 2011) to combine data on food availability and disturbance (Drost et al. 2001) and to study the capacity for geese under various


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levels of grazing and predation pressure (Stahl & Loonen 1998). The model is also relevant for the proposals of Van den Brink and Hacquebord in this core program as it can evaluate habitat use in relation to pollution and eutrophication. 2. Determine meta-population structure and linking genetic markers to fitness If we would be able to recognize colony specific genes, we could study the dynamics of these genes and genetic diversity over a period of colony establishment, rapid growth, stabilization and decline.

The usual pattern of population growth in a colony is a period of rapid growth short after its establishment based on local reproduction and return of female offspring to the natal colony. When geese become breeders, they are traditional and return to the same colony. With the high reproductive potential of geese, the new colonies reach saturation within a few decades. This colony saturation causes a decline in reproductive output with age after an initial increase due to gaining experience (Black et al. 2007).

In Kongsfjord, the colony established in1980. Since 1989, geese are ringed. In the early years reproductive output was high, but it turned to practically zero in the period 2000-2010. Despite this low local reproduction the number of nests kept increasing, suggesting influx from other colonies.

Molecular techniques may be applied to analyse contemporary patterns of divergence, population size and gene flow (Palsbøll et al. 2010). Until now, 651 individuals have been blood sampled for DNA, the oldest individual from 1989. A first attempt to study gene flow between barnacle goose populations has recently been made based on a genome wide set of 358 SNPs (Jonker 2012, 2011). With this proposal, we will expand on genome-wide SNP genotyping using RAD-tagged genomic sequencing (Peterson et al. 2012) which generates 1000s of SNP genotypes in each individual in order to improve gene flow estimates at different temporal and spatial scales. Moreover, we will also look for attributes of individual genes, which may correlate with fitness or pathogen resistance, or where there is a link with partner choice as found for the major histocompatibility complex (Richardson et al. 2005). In addition, we will estimate long-term trends in abundance from so-called skyline plots using the coalescent-based Bayesian approach implemented in the software BEAST (Drummond et al. 2007), as is planned in the core project headed by Palsbøll aimed at historic whale abundance. 3. Determine the ecological niche of three goose species to understand habitat segregation.

In the experimental set-up employed during the EU-program FRAGILE little attention was paid to niche differentiation among goose species, intra-specific competition, as well as species-specific differences in historical trends. In this study we want to highlight the importance of this information for understanding changes in the Arctic ecosystem and to enable realistic predictions for the future.

The oldest records of geese on Spitsbergen (Lövenskiold 1964) refer to large numbers of brent geese (Branta bernicla hrota) breeding along the West coast. During the last decades brent geese have become increasingly rare in these areas. Brent geese first changed area to East-Svalbard, where they were subject to heavy predation from the recovering polar bear population (Madsen et al. 1989). A later change to breeding grounds in Northwest Greenland, escaped this issue and the population has increased in numbers as noted on the Danish wintering grounds (Clausen et al. 2003). Brent geese are relatively more capable of foraging on salt sprayed vegetation compared to barnacle geese (Stahl et al. 2002).

Pink-footed geese (Anser brachyrhynchos) nests dispersed across the tundra but have always been reported as common (Strijbos 1957). They are still nesting across Svalbard and have increased in numbers despite active hunting and use of limited stop-over habitat during spring migration (Fox et al. 2010).

In contrast, barnacle geese (Branta leucopsis) used to be rare on Spitsbergen. However, the population grew when hunting was banned and protection measures were put in place on the wintering grounds. As a result their abundance has increased 100-fold over the last 70 years. This tremendous expansion lead to barnacle geese moving into areas formerly occupied by pink-footed and brent geese and barnacle geese now outnumber other goose species on the West coast of Spitsbergen. The nesting sites of barnacle geese changed from bird cliffs to small islands, which could accommodate the massive population expansion. However, increasing predation pressure by polar


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bears on these small islands might change the prospect of successful island breeding (Drent & Prop 2008).

Our present knowledge on niche differentiation among these species during the Arctic summer is based on a habitat-specific risk of predation and a species-specific difference in withstanding a predator. Far less is known about the ecological niche of the three above goose species under similar circumstances, their species-specific effect on the vegetation and species-specific immunology (see also item 4). In this project a common garden experiment with adults and goslings will assess these species-specific differences Experimental setup

Individuals from all three goose species are tested in captivity under similar conditions on Arctic tundra vegetation. Measurements on intake rate will be based on direct observations of bite size, peck speed and faeces production in combination with digestibility of food (Kuijper et al. 2009). Diet selection will be based on direct observation and analysis of faeces, where we will use microscopical inspection of epidermis fragments and molecular techniques (Stech et al. 2010, proposal Kruijer). Species specific critical lower biomass for intake will be derived from these measurements. Dominance interactions will be quantified in mixed flocks where different species compete for food. A test on salt sprayed vegetation provides species-specific effects on food intake (Stahl et al. 2002). Direct effects on the grazed plants will be examined by following the fate of grazed plants.

On a weekly basis body condition will be assessed as body mass and size. These measurements will be combined with immunological tests of blood samples (2-4 ml). The specific suite of tests is described in detail under item 4.

We will begin with three juveniles per species. The next year, we will adapt our experiment based on a power analysis of the results obtained during the first year. When relevant, the experiment will also include measurements on species-specific effects on the grazed vegetation. 4. Study differences in immunological parameters and pathology in relation to goose species and pollution level.

The Arctic has proven to be a relatively germ-free environment for raising offspring (Piersma 1997, Prop et al. in prep). Nevertheless, barnacle geese have proven to breed successfully at lower latitudes. At present barnacle geese are the fastest increasing breeding bird in the Netherlands and while breeding started in 1980, the population size of the Dutch breeding population of barnacle geese is now similar in size of the whole Svalbard population (more than 30.000 individuals). The ability of the barnacle goose to breed successfully in the Netherlands is especially striking because brent and pink-footed geese have not been recorded and, seem incapable of doing so. An important hypothesis emerging from the research conducted under the IPY-project BIRDHEALTH is that barnacle geese are capable of adapting to higher pathogen and parasite loads compared to the two other goose species. This may also explain the difference observed in captive breeding. Arctic species, such as the pink-footed and brent goose, rarely reproduce in captivity and are thus rare in wildfowl collections.

During the project BIRDHEALTH, we estimated a higher rate of survival among adult geese in pristine areas compared to human-affected areas on Spitsbergen (Prop et al., in prep). The higher survival rates were correlated with lower levels of natural antibodies as measured by the lytic performance of plasma on foreign (rabbit) red blood cells (Matson et al. 2005). This correlation is further explored in project Van den Brink. Methodology Blood samples taken from the experimental birds (above) or wild birds under comparable conditions (e.g. foraging in or outside of a polluted area) will serve as the basis for immunological tests. Microscopical inspection of blood smears will render a differential white blood cell count. In vitro assays will involve bactericidal activity of whole blood (Tieleman et al. 2005) and a hemolysis-hemagglutination test (Matson et al. 2005). Each individual will be challenged with sheep red blood cells (Deerenberg et al. 1997) and a PHA swelling test (Martin et al. 2006). External parasites are examined by the bag-ruffle method and visual inspection (Walther & Clayton 1997). Faeces will be collected to assess the rate of shedding of intestinal parasites (Dolnik & Loonen, 2006). At the end of the experiment, the geese will be terminated and given a full pathological examination. Biopsies will be taken for measurement of species specific levels of pollutants and stable isotopes.


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Societal and global significance The innovative aspect of this project lies in the integration of recent developed techniques and

insights to reveal detailed interactions within this Arctic terrestrial food web. All proposed methodology has been used within the research group. Geese in the Arctic have a direct effect on vegetation and predator populations and play a significant role in the arctic terrestrial ecosystem. Goose numbers have increased to a level where management plans are developed to reduce population sizes. In Europe the population reduction is motivated by damage to agriculture, whereas in North America the protection of the Arctic tundra is the main motivation. These management plans are likely to alter inter-specific competition. Understanding of historical patterns is needed to enable predictions concerning the consequences of human-induced drivers on the future scenarios of the Arctic terrestrial tundra ecosystem.

Literature cited Baveco, J. M., H. Kuipers, and B. A. Nolet. 2011. A large-scale multi-species spatial depletion model

for overwintering waterfowl. Ecological Modelling 222:3773-3784. Black JM, Prop J & Larsson K (2007) Wild goose dilemmas. Branta Press, Groningen. Clausen P, Green M & Alerstam T (2003) Energy limitations for spring migration and breeding: the

case of brent geese Branta bernicla tracked by satellite telemetry to Svalbard and Greenland. Oikos 103: 426-445.

Deerenberg C, Apanius V, Daan S & Bos N (1997) Reproductive effort decreases antibody responsiveness. Proc. R. Soc. Lond. B 264: 1021-1029.

Dolnik OV & Loonen MJJE (2006). First finding of Tyzerria parvula (Kotlàn, 1993) Klimeš, 1963 (Protozoa: Coccidiida) in Barnacle Geese (Branta leucopsis Bechstein, 1803) on Spitsbergen. Zoosystematica Rossica 15: 214.

Drent RH & Prop J (2008) Barnacle goose Branta leucopsis survey on Nordenskiöldkysten, west Spitsbergen 1975–2007: breeding in relation to carrying capacity and predator impact. Circumpolar Studies 4: 59-83.

Drost A, Kruckenberg H & Loonen MJJE (2001) Untersuchungen zur Störungsempfindlichkeit arktischer Nonnengänse während der Brut- und Mauserzeit. Vogelkdl Ber Niedersachs 33:137-142.

Drummond AJ & Rambaut A (2007) "BEAST: Bayesian evolutionary analysis by sampling trees." BMC Evolutionary Biology 7: 214.

Ebbinge BS & Spaans B (2002) How do brent geese (Branta b. bernicla) cope with evil? Complex relationships between predator and prey. Journal für Ornithologie 143: 33-42.

Fox AD, Ebbinge BS, Mitchell C, Heinicke T, Aarvak T, Colhoun K, Clausen P, Dereliev S, Faragó S, Koffijberg K, Kruckenberg H, Loonen MJJE, Madsen J, Mooij J, Musil P, Nilsson L, Pihl S, & Van der Jeugd H (2010) Current estimates of goose population sizes in western Europe, a gap analysis and an assessment of trends. Ornis svecica 20: 115-127.

Jefferies RL, Rockwell RF Abraham KF (2004) Agricultural food subsidies, migratory connectivity and large-scale disturbance in arctic coastal systems: a case study. Integr. Comp. Biol. 44: 130-139.

Jensen RA, Madsen J, O’Connel M, Wisz MS, Tømmervik H & Mehlum F (2007) Prediction of the distribution of Arctic-nesting pink-footed geese under a warmer climate scenario. Global Change Biology 14: 1-10.

Jonker RM, Zhang Q, van Hooft P, Loonen MJJE, van der Jeugd HP, Crooijmans RPMA, Groenen MAM, Prins HHT & Kraus RHS (2012) The development of a genome wide SNP set for the Barnacle Goose Branta leucopsis. PLoS ONE (accepted).

Jonker RM, Kraus RHS, Zhang Q, van Hooft P, Larsson K, van der Jeugd HP, Kurvers RHJM, van Wieren SE, Loonen MJJE, Crooijmans RPMA, Ydenberg RC, Groenen MAM & Prins HHT (2011) Genetic consequences of breaking migratory decisions in barnacle geese. In: Jonker R.M. Revolutionary non-migrants. Ph. D. Thesis Wageningen University.

Kotanen PM & Jefferies RL (1997) Long-term destruction of sub-arctic wetland vegetation by lesser snow geese. Ecoscience 4:179-182.

Kuijper DPJ, Ubels R & Loonen MJJE (2009) Density-dependent switches in diet: a likely mechanism for negative feedbacks on goose population increase? Polar Biology 32: 1789-1803.


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Loonen MJJE (1997) Goose breeding ecology: overcoming successive hurdles to raise goslings. Ph. D Thesis University of Groningen.

Loonen MJJE, Tombre IM & Mehlum F (1998) Development of an arctic barnacle goose colony: interactions between density and predation. Norsk Polarinstitutt Skrifter 200: 67-79.

Loonen MJJE & Solheim B (1998) Does arctic vegetation change when grazed by barnacle geese? A pilot study. Norsk Polarinstitutt Skrifter 200: 99-103.

Løvenskiold, HL (1963) Avifauna svalbardensis. Norsk Polarinstitutt Skrifter 129. Madsen J & Bregnballe T & Mehlum F (1989) Study of the breeding ecology and behaviour of the

Svalbard population of Light-bellied Brent Goose Branta bernicla hrota. Polar Research 7: 1-42.

Martin II LB, Han P, Lewittes J, Kuhlman JR, Klasing KC, Wikelski M (2006) Phytohemagglutinin-induced skin swelling in birds: histological support for a classic imunnoecological technique. Functional Ecology 20: 290-299.

Matson KD, Rickleffs RE & Klasing KE (2005) A hemolysis-hemagglutination assay for characterizing constitutive innate humoral immunity in wild and domestic birds. Developmental and Comparative Immunology 29: 275-286.

O'Connell MJ, Huiskes AHL, Loonen MJJE, Madsen J, Klaassen M & Rounsevell M (2006) Developing an integrated approach to understanding the effects of climate change and other environmental alterations at a flyway level. Waterbirds around the world. Eds. G.C. Boere, C.A. Galbraith & D.A. Stroud. The Stationary Office, Edinburgh, Uk. pp. 385-397.

Palsbøll PJ, Peery MZ, & Bérubé M (2010). Detecting populations in the “ambiguous” zone: kinship-based estimation of population structure at low genetic divergence. Gaggiotti OE (ed): Special Issue: advances in the analysis of spatial genetic data. Molecular Ecology and Resources 10: 797-805.

Peterson BK, Weber JN, Kay EH, Fisher HS & Hoekstra HE (2012) Double Digest RADseq: An Inexpensive Method for De Novo SNP Discovery and Genotyping in Model and Non-Model Species. PLoS ONE 7(5): e37135. doi:10.1371/journal.pone.0037135.

Piersma T (1997) Do global patterns of habitat usen and migration strategies co-evolve with relative investments in immunocompetence due to spatial variation in parasite pressure? Oikos 80: 623-631.

Prop J, Griffin LR, van der Jeugd HP & Loonen MJJE (in prep) Evolutionary benefits of breeding in the Arctic.

Richardson DS, Komdeur J, Burke T & von Schantz, T (2005) MHC-based patterns of social and extra-pair mate choice in the Seychelles warbler. Proc. R. Soc. B 272: 759-767.

Sjögersten S, van der Wal R, Loonen MJJE &, Woodin SJ (2010a) Recovery of ecosystem carbon fluxes and storage from herbivory. Biochemistry 106: 357-370.

Sjögersten S, Kuijper DPJ, van der Wal R., Loonen MJJE, Huiskes AHL. & Woodin SJ (2010b) Nitrogen transfer between herbivores and their forage species. Polar Biology 33: 1195-1203.

Speed JPM, Woodin SJ, Tømmervik H, Tansdorf P & van der Wal R (2009) Predicting habitat utilization and extent of ecosystem disturbance by an increasing herbivore population. Ecosystems 12: 349-359.

Stahl J & Loonen MJJE (1998) The effects of predation risk on site selection of barnacle geese during brood-rearing. Norsk Polarinstitutt Skrifter 200: 91-98.

Stech M, Kolvoort E, Loonen MJJE, Vrieling K & Kruijer JD (2010) Bryophyte DNA sequences from faeces of an arctic herbivore, barnacle goose (Branta leucopsis). Molecular Ecology Resources 11: 404-408.

Strijbos, JP (1957) Svalbard. Zwerftocht langs de koele stranden van Spitsbergen. Veen, A’dam. Tieleman BI, Williams JB, Ricklefs RE & Klasing KC (2005) Constitutive innate immunity is a

component of the pace-in-life syndrome in tropical birds. Proc. R. Soc. B 1573: 1715-1720. Van der Jeugd HP, Eichhorn G, Litvin KE, Stahl J, Larsson K, van der Graaf AJ & Drent RH (2009)

Keeping up with early springs: rapid range expansion in an avian herbivore incurs a mismatch between reproductive timing and food supply. Global Change Biology 15: 1057-1071.

Van der Wal R, van Lieshout SMJ, & Loonen MJJE (2001) Herbivore impact on moss depth, soil temperature and arctic plant growth. Polar Biology 24: 29-32.

Van Geest GJ, Hessen DO, Spierenburg P, Dahl-Hansen GAP, Christensen G, Faerovig PJ, Brehm M, Loonen MJJE & Van Donk E (2007) Goose-mediated nutrient enrichment and planktonic grazer control in arctic freshwater ponds. Oecologia 153: 653-662.


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Walther BA & Clayton DA 1997. Dust ruffling: a simple method for quantifying ectoparasite loads of live birds. J.. Field Ornithol.. 68:509-518.

Zielke M, Loonen MJJE & Solheim B (2004) Response of nitrogen fixation and biomass productivity on long-term grazing and fertilization by Barnacle geese (Branta leucposis) in high arctic tundra vegetation. In: Zielke M. Diversity and nitrogen fixation activity of cyanobacterial communities in terrestrial arctic ecosystems. Doctoral thesis, University of Tromsø, Norway.

10. Timetable of the project

The project has three seasons of field work. The first season is for preparation of the experiment, and finalising the measurements for the spatial distribution model. In year 2 and 3, the experiment for objective 3 will be conducted. The first period of data analysis is based on existing data sets (objective 1). The second period is following laboratory analysis for objective 2. The third period is focussing on the analysis (objective 4) of the first field experiment (objective 3). The fourth period of analysis is a final preparation for the final writing session. Manuscript writing is also envisaged in the first year to gain experience.

11. Fieldwork 11a. What is/ are the proposed fieldwork location(s)? (region, town, station)

Svalbard, Spitsbergen, Ny-Ålesund, Netherlands Arctic Station in close cooperation with the Norwegian Polar Institute 11b. What is /are the proposed period(s) of fieldwork?

Start Date (mm, yyyy) End Date (mm, yyyy) Duration (weeks)

07,2013 08,2013 6 weeks

06, 2014 08. 2014 11 weeks

06, 2015 08, 2015 11 weeks

11c. What is the proposed way of transportation to the fieldwork location(s)?

Airplane 11d. Will you be making use of the NIOZ-MRF pool of equipment?

Ο Yes X No 11e. Has been complied with the law and legal requirements with respect to the proposed research, such as ‘DNA-recombinant legislation’, ‘Code Openness Animal Experiments’, and/or ‘Code of conduct on Biosecurity’, import of samples

X Yes Ο No


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12. Data management Please provide information concerning filing of your data 12a. What kind of data will you collect with this project?

Most data will be linked to marked individuals: life history parameters, geo-referenced spatial distributions, genome sequence data, immunological tests, health status, behaviour. General data will involve population estimates, vegetation and assessments of the environment. 12b How and when do you intend to give public access to your data

For all data we will describe and publish meta data on the internet and submit the meta data to international collections. All data will be organized so it can be stored at datacenters specifically equipped for that type of data. The original data will only become available for the public after scientific publication. The main focus of this program will be to produce scientific publications based on the available data in peer-reviewed journals (preferably open-access). After the project, a popular text book is planned on the terrestrial ecosystem. 12c. What is the long term (in terms of decades) data archiving plan for this proposal

That depends on the state of the art. At present, two data management projects would be used. The Polar data center of NWO hosted by NIOZ Texel and the Research in Svalbard (RiS) database, hosted by the Svalbard Science Forum

13. (Inter)national collaboration Is this proposal part of an (inter)national collaboration? If ‘yes´, with whom?

This proposal links directly to two international projects: FRAGILE (EU-FP5 2003-2005) and BIRDHEALTH (IPY-NL 2007-2009). Although both projects have ended, the international collaborative structure is still present and new papers are appearing as spin-off of these projects. The goose-related studies of the Netherlands Arctic Station in Ny-Ålesund have always been in close collaboration between Dutch, Norwegian and British partners. The long term ecological data have been used as input in the ARCTIC WOLVES project (IPY-Canada) and for scientific presentations in NySMAC, the CBIRD group of CAFF, the IWRB-GSG goose specialist group and to obtain observer status in the EU-supported INTERACT consortium. Name Affiliation Expertise History Dr J. Madsen Arhus University, Denmark pink footed goose FRAGILE,IWRB-GSG,

BIRDHEALTH

Dr S.J. Woodin Univ. Aberdeen, UK vegetation, N-cycle FRAGILE

Dr R. van der Wal Univ. Aberdeen, UK vegetation, decomposition

FRAGILE

Dr M. Klaassen Deakin Universituy, Australia

migration, influenza FRAGILE,BIRDHEALTH

Dr A.D. Fox Arhus University, Denmark goose ecology FRAGILE,IWRB-GSG

Dr P. Clausen Arhus University, Denmark while-bellied brent geese IWRB-GSG

Dr H. Kruckenberg Univ. Osnabrück, Germany Russian geese IWRB-GSG

Dr. R.F. Rockwell Nat. Museeum, New York, USA

lesser snow geese ARCTIC WOLVES

Dr G. Gauthier Univ. Lavalle, Quebec, CA greater snow geese ARCTIC WOLVES

Dr K. Abraham Trent University, Peterborough, CA

goose management ARCTIC WOLVES

Dr G. Gilchrist Environment Canada, Ottawa

common eider ecology and diseases

CBIRD-CAFF

Dr L. Griffin WWT, Slimbridge, UK ringing scheme FRAGILE,IWRB-GSG

Dr. H.H.T. Prins Wageningen University ecology IWRB-GSG

FRAGILE: http://cordis.europa.eu/search/index.cfm?fuseaction=proj.document&PJ_RCN=6158452 BIRDHEALTH: http://www.birdhealth.nl ARCTIC WOLVES: http://www.cen.ulaval.ca/arcticwolves NySMAC: http://nysmac.npolar.no/ CBIRD-CAFF: http://caff.is/cbird IWRB-GSG: http://www.geese.org/gsg/ INTERACT: http://www.eu-interact.org


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14. Finances 14a. Requested budget:

Year 1 Year 2 Year 3 Year 4

Personnel (mm) Research costs (k€) Equipment 8000 14500

Consumables*,† 10000 10000 10000 2400 Fieldwork/Travel† 4000 6800 6800 Bench fee 5000 *Max. k€10 per year †Max. k€40 voor een postdoc en k€50 voor een AiO. 14b. Specification of the requested funds:

Equipment: Behavioural registration (3 sets) € 4500; Geese holding facility and catching equipment € 4000;

Freezer storage € 8000; Safety equipment fieldwork € 1500; Dataloggers € 2000; Water resistant field data recorder € 2500 Total: € 22,500

Consumables: Molecular analysis per sampled individual: € 60 (€50 per sample for RADseq and €10 for DNA extraction and qPCR DNA quality check), 400 samples: € 24,000 Immunological tests per sampled individual: € 30 (based on earlier BIRDHEALTH project), 200 samples: € 6,000 Vegetation quality analysis (nitrogen, acid detergent fiber) 48 samples x € 50 = € 2400 Total € 32,400

Fieldwork/Travel: Travel and hotel to Ny-Alesund: €1200,- board: €70,- per day, fieldwork year 1,2,3= 40,80,80 days. Total € 17,600

14c. Financial assistance from other sources Please describe any financial assistance for this research proposal from other sources

Student projects during field work and laboratory analysis will be supported by the Willem Barentsz Foundation and by sponsoring. Lodging in the Netherlands Arctic Station and field work assistance by the thesis supervisor will be paid by the governmental grant for the Arctic station..

No signatures required for electronic submission

.

..1. Novelty/originality and innovation

..2. Scientific quality of the proposal

..3. Qualification of the applying research group

Subsidie / Subsidy : Nieuw Nederlands Polair Programma 2012 ALW

Projectnummer / Project number : ALWNNPP2012-1-12

Hoofdaanvrager / Main applicant : Dr. M.J.J.E. Loonen

Projecttitel / Project title : From historical data to a prediction of the future forgeese on arctic tundra?

Adviseur: 1

Confidential

RESEARCH PROPOSAL(You may extend the form to have it fit your comments, a typical review has 2 or 3 pages)

Judgement:

This study combines four different objectives covering the habitat use of geese, the genetic structure of apopulation with a known history, the niche differentiation of three goose species breeding in the Spitsbergen andthe immunological status of breeding barnacle geese in the arctic vs. temperate regions. The combination of suchdisparate approaches to study and predict future demography is particularly original, and it will lead to veryexciting results if the project team can pull all aspects together.

a. Scientific approach and methodologyb. Clarity and substance of objectivesc. Impact: potential extension/deepening of knowledged. Improvement and reinforcement of (inter)national scientific cooperation

Judgement:

Approach/MethodologyEach of the four aspects of the project seem to be sound. However, a weakness of this application is that it is notclear how the four approaches will be brought together to achieve the ultimate aim of predicting future status ofgeese in the arctic tundra.

Clarity/Substance of objectivesEach of the four objectives is clearly presented. However, how they relate together and how they will beintegrated to predict the future of geese on the arctic tundra is less clear.

ImpactGeese are key species in the arctic tundra and can have severe impacts on agricultural systems in their winteringand migratory grounds in northern Europe. Understanding their demography and predicting future trends in goosepopulations (of all species) is thus very important for future management of agricultural systems and remote arcticregions. This project will go some way to achieving this understanding. Each of the four objectives should lead tohigh quality peer-reviewed publications, and a PhD candidate will be educated in a great atmosphere between thedifferent aspects of the project, potentially with collaboration within the broader core program. However, theproject team will face a challenge in drawing the four objectives together to reach a prediction of the future ofgeese on the tundra.

CooperationThe project partners are from three institutions in the Netherlands with one partner from Norway, so this projectinvolves some but not much cooperation at the national and international level. However the research group hasa history of recent collaboration with partners in many (six) countries and if the project can build on these links inaddition it will serve to strengthen international cooperation.

Judgement:

Dossier nummer / File number: ALWNNPP2012-1-12

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4.

4.a. Does the proposal fit in the core programme

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The research group at the Arctic centre and the Netherlands research station at Ny Alesund is an internationallyrecognised hotspot for goose research. This is documented by the publication and achievement list and research/project history of the group. The research group is thus very well qualified for the ambitious aim of the project.

Core programmeEach proposal forms a contribution to a core programme. The proposals are supposed to be independent,though they need to fit in the core programme. The abstracts of all proposals of the core programme areincluded.

Judgement:

The keystone role of geese in the arctic ecosystem, and the relationship with humans means that this project fitsvery well into the core program and it seems that very close links could potentially be formed with other terrestrialcomponents within the program. Links with marine parts of the program are more tenuous however.

FINAL ASSESSMENT SCIENTIFIC QUALITY:

How do you value this proposal regarding the above mentioned qualifications? Please substantiate yourqualification in one or two concluding sentences:

- Excellent- Very good- Good- Reasonable- Poor

Concluding remarks:

Very good

If the project team are able to integrate the four aspects of this project it will produce important results for thesekey species in both the arctic and the agricultural systems in Northern Europe


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4.

Adviseur: 2

Confidential


Judgement:

Highly original and novel as it integrates key aspects of complementary disciplines in biology. Firstly, thedevelopment of a spatially explicit goose distribution model combining data on plants and disturbance is highlyrelevant for ecologist trying to anticipate the consequences of global changes on the structure and functioning ofthe arctic terrestrial ecosystem. Secondly, the use of molecular techniques to study gene flow at various temporaland spatial scale is original and relevant in the context of the actual shifts in arctic species distribution. Finally, theproject is investigating poorly known aspects of arctic species interactions (ecological niche; pathogen/parasiteseffects).


Judgement:

Overall assessment - HIGH QUALITY - very few research team in the world could potentially reach these originalobjectives using the proposed methodology.

a. The scientific approach and methodology: convincing and novel. Considering the quality of the experts involvedin the project as well as the qualifications of the collaborators, I anticipate no major methodological issues. Notethat the project seems ambitious for a PhD student as it is highly integrative and multidisciplinary - the use ofmultiple lab and modelling techniques, sources of data as well as arctic field experiment is a major strength of theproposal but may represent a real challenge for a PhD candidate.

b. In general, the four main objectives are relatively well presented and highly relevant. I found novel andimportant ecological aspects in all four objectives. However, the integration of the different objectives is not alwaysobvious in the proposal and the addition of theoretical framework/working hypotheses for each objective wouldhave improved the proposal.

c. Impact - The project will certainly generate extremely relevant and new knowledge in the field of biology. Isuspect that objectives 1, 2 and 4 could have the stronger potential impact if they are reached.

d. The project leader has already developed a very strong international network and the collaborative structure willlikely contribute to the success of the proposed project. The project fit well with international research initiativesdeveloped recently and this will very likely strengthen the linkages established during the International PolarYear.

Judgement:

This is one of the best research group in the world investigating arctic wildlife ecology. The quality and quantity oftheir scientific contributions are outstanding considering the challenges associated to long-term ecologicalmonitoring in the rapidly changing arctic ecosystem. The group perform highly relevant and innovative researchprojects having strong international impacts. The team is highly interdisciplinary and can tackle challenging andurgent research questions associated to the impact of global changes on polar regions. The team is in anexcellent position to make substantial scientific contribution in the short-term.

Core programmeEach proposal forms a contribution to a core programme. The proposals are supposed to be independent,


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....

though they need to fit in the core programme. The abstracts of all proposals of the core programme areincluded.

Judgement:

Yes. The project will generate important interdisciplinary knowledge that strongly fit with the core program of theArctic Centre. The multi-faceted of the project (including immunology, field experiment, population genetic) willplace he selected PhD candidate in an excellent position to integrate knowledge produced by other teammembers and to generate new data relevant to other proposals (including other proposals on arctic herbivores,arctic plants, arctic contaminants and species interactions).




Concluding remarks:

Excellent. This project is innovative, interdisciplinary and tackle important research questions.The research will be performed using novel techniques, short-term field experiments as well as existing, rare andunique long-term data. This project will likely generate very high-quality scientific papers with substantial impact inthe international scientific community.


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Adviseur: 3

Confidential


Judgement:

I paid attention to the PhD proposal presented by Prof. M.J.J.E. Loonen on the dynamic of an arctic goosespecies. This multi-disciplinary research proposal is very appealing in the sense that it would combine informationon genetic, physiology (immunology), foraging behaviour including diet, competition and habitat selection as wellas ecosystem functioning and would make use of long-term dataset and a field experiment. All these approachesare to my point of view totally justified to understand past and present abundance of geese and assess theimportance of direct and indirect impact of climate change and anthropogenic disturbances to be able to predictfuture population trends and the use by geese of the tundra.


Judgement:

a. Scientific approach and methodologyThe scientific approaches in the four proposed chapters are accurate and will address the questions raised here.For each objective, I provide several comments

Objective 1 focuses on trophic interactions. The authors propose to adopt a modelling approach based onenergetic requirements and functional responses of geese (Baveco et al. 2011). Although, this approach is verydetailed and would establish the carrying capacity for geese breeding in the Arctic, it requires a lot of parametersthat would mostly come from literature (except the functional response that will be studied in chapter 3). I questionhow disturbance, predation risk, allochtonous subsidies (body reserves, lipids that contribute to egg formation ingeese, see Gauthier et al. 2003 Ecology on Snow geese) and predation (on eggs, juveniles or even adults) wouldbe incorporated in such models. Although, all this could probably be incorporated in such energetic modellingframework, I would suggest the use of food-web models based on mass balance that also depend on functionalresponses (i.e., DeRuiters et al. 1995 Science). It could be an alternative maybe less complicated that would havethe advantage to incorporate directly the different trophic levels of this ecosystem. This modelling approach is alsoa conceptual framework (ecosystem functioning, meta ecosystems; Loreau 2011 Princeton) that would particularlybe relevant for this project and for the whole core program that attend to describe past present and future dynamicof marine, terrestrial and freshwater ecosystems.

Objective 2. Dispersal is a key factor driving populations dynamic and still poorly understood and using moleculartechniques on an existing long-term dataset is very exciting promising. As a non-geneticist, I am not able to judgethe methodological approach presented here but the fact that preliminary analyses were already conducted isencouraging.

Objective 3. Identifying functional responses of geese and investigate potential competition by conducting acommon garden experiment in the arctic is just lacking and it would definitely tease apart potential confoundingeffects that can emerge when the niche is established on habitat selection only. I would tend to combine thisexperiment with the project of Prof. Kondeur (project 06 of the core programme) and investigate stress (probablyglucocorticoids CORT on droppings I guess) of geese under the different treatment. This would give some insighton the effect of the feeding treatment not only by measuring the body condition (based on biometricmeasurements) but also by measuring its effect on the physiology and possible consequences of chronic stress.

Objective 4. Understanding evolutionary process of bird migration is still under investigated and the influence ofparasites is particularly relevant. The idea that the immune system of barnacle geese is more adaptive than theother species is intriguing and it should deserve attention. It is not explicitly stated if such project would beconducted in the Dutch geese populations. It would be very interesting in a perspective to explain geese


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4.


.

....

migration. I also wonder if the barnacle geese that breed in the Netherlands could (partly) be issued from feralindividuals released into the wild ? This might explain their capability to mount stronger immune responses?

b. Clarity and substance of objectivesEach objectives are clear, straightforward and promising.

c. Impact: potential extension/deepening of knowledgeAs stated earlier, I would tend to recommend to explore how food-web models could be another avenue todescribe trophic interactions and how perturbations (through simulations of different scenarios of resourceabundance or with the addition of other species or subsidies, see Leroux and Loreau 2008 Ecol. Lett. could modifypatterns of ecosystem functioning and in turn affect geese populations). This might be anecdotic, but the authorsshould mentioned in their proposal that Svalbard terrestrial ecosystem is unique compare to other places in thesense that there is no lemming which greatly simplify the food web and strengthen the fact that geese are keyplayers in the tundra. The interaction between reindeer and geese should also be of interest. Also, I found that thisproject mostly emphasized on the interaction between plant and herbivores but predation can also be an importantdriver in tundra ecosystems (e.g. Gilg et al. 2003 Science; Legagneux et al. 2012 Ecology).

d. Improvement and reinforcement of (inter)national scientific cooperationThis project is based on the collaboration between scientists of different horizons that would work together withthe PhD candidate. For each objective, the scientific cooperation seems relevant. I would recommend discussingwith ecosystem modellers and theoreticians that may have a different approach to address the predator-preyinteraction study. For the other objectives, the cooperation seems adequate.

Judgement:

All the partners involved in this project are recognised international scientists. The research group is solid andwould be able to address the objectives as long as the PhD candidate is carefully selected. The PhD candidatewill indeed have to play with many concepts from various ecological disciplines (ranged from micro to macroscales) and interact with scientists from diverse origins, which makes this project very challenging and sointeresting.

Core programmeEach proposal forms a contribution to a core programme. The proposals are supposed to be independent,though they need to fit in the core programme. The abstracts of all proposals of the core programme areincluded.

Judgement:

-




Concluding remarks:

This proposal is ambitious and achievable because the expertise of the supervisors, mentors and their experienceof the field site is solid enough to conduct such a multidisciplinary study. For all the reasons mentioned above, Ievaluate this proposal as VERY GOOD


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NWO

REBUTTAL Rebuttal form ALW Open programme (max. 2 pages, including figures, font Arial 10 point) In pursuit of fair comparison between projects, rebuttals longer than two pages can not be accepted Main applicant : Dr. M.J.J.E. Loonen Title : From historical data to a prediction of the future for geese on arctic tundra? File number : ALWNNPP2012-1-12

We are pleased with the reviewers rating our proposal once excellent and twice very good. The remarks made by the latter two reviewers relate to the last sentence in our summary “All elements combined should enable the prediction of future scenarios of anthropogenic impact on species interaction within the food web.” This sentence clearly defines our ultimate goal but not the final product of this proposal. In our introduction to the description of the proposed research we state: “The project will provide building blocks for modelling future scenarios for goose demography on the Arctic tundra.” The notion of building blocks is best suited for the final goal of this project. Our prior involvement in the EU-project FRAGILE Fragility of Arctic Goose habitat: impacts of land use, conservation and elevated temperatures and the IPY-project Arctic Wolves Arctic wildlife observatories linking vulnerable ecosystems has learned us, that we can contribute to the ultimate goal, with elements which fit in our expertise and our long-term experience collecting field data. In the proposed project, we are focussing on four understudied objectives to understand historical changes in goose distribution. Our long-term study has identified these objectives as potential crucial for a better understanding of the food web and a prediction of the future for geese on arctic tundra. All reviewers recognize this approach as “highly original and novel” (ref 2), “particular original, and will lead to exciting results” (ref 1) and “totally justified to understand past and present abundance of geese and assess the importance of direct and indirect impact of climate change and anthropogenic disturbances to be able to predict future” (ref 3). Referee 1 describes as weakness: “how the four approaches will be brought together”. Our focus on these four objectives does not mean that we consider these as the only elements to contribute to understanding of future scenarios. To understand the food web there are many more relevant elements of which many are already under study by us. We have focussed on objectives which are within reach, novel and relevant and which can be studied at the pre-defined study site (pre-defined by the call) by one Ph.D. working together with other projects in the core program. Objective 1 (Develop a spatial explicit distribution model for grazing pressure based on tropic interactions and anthropogenic driver) is clearly one type of a modelling approach to integrate existing local data from the long-term study and other projects in the core program. Objective 2 (Determine meta-population structure and linking genetic markers to fitness) will expand the data studied under objective 1 and 4 with a new perspective based on molecular techniques. Objective 3 (Determine the ecological niche of three goose species to understand habitat segregation)and 4 (Study differences in immunological parameters and pathology in relation to goose species and pollution) focus on important elements which have received little attention so far and have the potential to develop into a key factor for objective 1 and the prediction of future scenarios for geese on Arctic tundra. We have focussed on four different objectives to optimally integrate this project with several other projects in the core program (Hacquebord, Van den Brink, Palsbøl, Kruijer and Komdeur) both in scientific and logistic terms. The links between the objectives and the other projects will certainly improve while the program is in progress based on new insights being gained. Referee 3 raised an interesting point to include food web models as alternative avenue to describe trophic interaction. We did consider this while designing the project but have chosen to limit the type of model to a spatial explicit distribution model, despite some limitations in scaling all actors and processes into one concept. We consider these models easier to test in the local situation, better

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linked with our spatial and temporal explicit long term dataset and closer to an integration with other (mapping) projects in the core program. The causes of annual fluctuations in abundance of various predatory species and even geese are not within our study area. This is also the main reason why we might seem to focus in our experiments on the interaction between plants and herbivores. We don’t agree with referee 3 that we miss on acknowledging predation as a driving force with this approach. In comparison with Bylot Island (Legagneux et al. 2012), our food web has no lemmings. However most important is that 70-100% of the annual primary production is removed by geese in their brood rearing area, compared to less than 10% on Bylot Island. This comparison clearly shows that bottom-up forces might be more important in our food web. Within our study area, we can test this experimentally. Top down control has also our full attention. With emerging presence of polar bears (Drent & Prop 2008) and effects of arctic foxes on body condition of the geese (Loonen et al. 1998, Stahl & Loonen 1998), we think that most value in understanding this trophic level is with the simple continuation of our long-term observations and not with the development of new experiments. Finally referee 3 advices to combine our experiment under objective 3 with the project of Komdeur. This was our intention from the moment we initiated the core program and both projects. Furthermore the Dutch goose populations are part of another Ph.D. project currently running at the Arctic Centre. So both remarks I consider relevant but already incorporated. We are very sorry that our proposal has generated some remarks from referee 1 and 3 concerning the scientific approach and methodology. Despite their very positive and stimulating remarks on the novelty/originality, these concerns shifted their overall scientific assessment from excellent to very good. We were aware of such a danger when combining four objectives and a multi-faceted approach in order to optimize the links between projects in the core program and to stimulate multi-disciplinary arctic research. We hope to have explained our considerations better in this rebuttal. The present project is based on our long-term study and presence in Ny-Ålesund and will be the first Ph.D. allotted by the Netherlands Polar Program to Loonen as principal investigator.. All three referees are exceptional positive about the qualifications of the applying research group (“very well qualified”, “one of the best research groups in the world investigating arctic wildlife ecology”, “recognised international scientists”). We thank the referees in their trust and hope that these statements will help the project to become funded. Groningen 12 Oct 2012 Maarten Loonen Drent RH & Prop J (2008) Barnacle goose Branta leucopsis survey on Nordenskiöldkysten, west

Spitsbergen 1975-2007: breeding in relation to carrying capacity and predator impact. Circumpolar Studies 4: 59-83.

Legagneux P et al. (2012) Disentangling tropic relationships in a High Arctic tundra ecosystem through food web modelling. Ecology 93: 1707-1716.

Loonen, MJJE, Tombre, IM & Mehlum, F (1998) Development of an arctic barnacle goose colony: interactions between density and predation. Norsk Polarinstitutt Skrifter 200: 67-79.

Stahl, J & Loonen, MJJE (1998) The effects of predation risk on site selection of barnacle geese during brood-rearing. Norsk Polarinstitutt Skrifter 200: 91-98.

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Ph.D. position for a biologist at the Arctic Centre of the ... · The PhD candidate will be investigating dynamics and structure of an Arctic terrestrial food web, with geese as the