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.
2.
3.
4.
5.
6.
A motivation letter.
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).
A certified copy of your college transcript (‘doctorandus’ or MA degree, or an
equivalent), as well as a list of examination results.
Email- and postal addresses of two academic referees.
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
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 interdisciplinary 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 fulltime 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
The maximum length of a proposal is 11 pages
1. Details of proposal
Theme:
Title:
Ο Ice, climate and sea level
Ο Polar oceans
X Polar ecosystem
Ο Humanities and changes in polar regions
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:
Institution:
Position:
Dr. Maarten J.J.E. Loonen
University of Groningen, Arctic Centre
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:
Tel:
Email:
Prof. Dr. L. Hacquebord
050-363 6832
[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
Page 1 of 11
New Netherlands Polar Programme
Proposal form 2012
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: 11171122.
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
Page 2 of 11
New Netherlands Polar Programme
Proposal form 2012
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
PhD, TBA
ecologist
Prof Dr P Palsbøl
genetic markers
population size
life history
genomics
polar
phytoplankton
goose ecology
Prof Dr J
Komdeur
Prof Dr AGJ
Buma
Dr. J Prop
Dr. BA Nolet
Prof Dr A Gröne
modeling
herbivore
behaviour
pathology
Dr. GW
Gabrielsen
Dr. A.J.M.
Scheepstra
arctic
ecotoxicology
coordination /
organization
University of
Groningen, AC
University of
Groningen, AC
University of
Groningen, CEES
University of
Groningen, CEES
University of
Groningen, ESRIG
University of
Groningen, AC
NIOO, Wageningen
University Utrecht,
Veterinary Medicine
Norwegian Polar
Institute
Willem Barentsz
Polar Institute
thesis supervisor, daily
guidance
project researcher
40
advisor, laboratory facilities
2
advisor, laboratory facilities
2
laboratory facilities, training
p.m.
advisor
4
advisor
p.m.
biopsies, training
p.m.
advisor, link to NPI
datasets and researchers
organizing workshops,
overarching PhD and
Postdoc projects
p.m.
7. Summary for the general public (in Dutch, max. 100 words)
6
2
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
Page 3 of 11
New Netherlands Polar Programme
Proposal form 2012
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: 17891803.
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
Page 4 of 11
New Netherlands Polar Programme
Proposal form 2012
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 socalled 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 EastSvalbard, 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
Page 5 of 11
New Netherlands Polar Programme
Proposal form 2012
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 pinkfooted 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 hemolysishemagglutination 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.
Page 6 of 11
New Netherlands Polar Programme
Proposal form 2012
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 interspecific 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.
Page 7 of 11
New Netherlands Polar Programme
Proposal form 2012
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) Phytohemagglutinininduced 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: kinshipbased 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: 623631.
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.
Page 8 of 11
New Netherlands Polar Programme
Proposal form 2012
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
Page 9 of 11
New Netherlands Polar Programme
Proposal form 2012
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
FRAGILE,IWRB-GSG,
Arhus University, Denmark
pink footed goose
Dr J. Madsen
Dr S.J. Woodin
Dr R. van der Wal
Univ. Aberdeen, UK
Univ. Aberdeen, UK
Dr M. Klaassen
Deakin Universituy,
Australia
Arhus University, Denmark
Arhus University, Denmark
Univ. Osnabrück, Germany
Nat. Museeum, New York,
USA
Dr A.D. Fox
Dr P. Clausen
Dr H. Kruckenberg
Dr. R.F. Rockwell
vegetation, N-cycle
vegetation,
decomposition
migration, influenza
goose ecology
while-bellied brent geese
Russian geese
lesser snow geese
Dr G. Gauthier
Dr K. Abraham
BIRDHEALTH
FRAGILE
FRAGILE
FRAGILE,BIRDHEALTH
FRAGILE,IWRB-GSG
IWRB-GSG
IWRB-GSG
ARCTIC WOLVES
ARCTIC WOLVES
Univ. Lavalle, Quebec, CA
greater snow geese
ARCTIC WOLVES
Trent University,
goose management
Peterborough, CA
CBIRD-CAFF
Environment Canada,
common eider ecology
Dr G. Gilchrist
Ottawa
and diseases
FRAGILE,IWRB-GSG
WWT, Slimbridge, UK
ringing scheme
Dr L. Griffin
IWRB-GSG
Wageningen University
ecology
Dr. H.H.T. Prins
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
Page 10 of 11
New Netherlands Polar Programme
Proposal form 2012
14. Finances
14a. Requested budget:
Year 1
Year 2
Personnel (mm)
Research costs (k€)
Equipment
8000
14500
Consumables*,†
10000
10000
Fieldwork/Travel†
4000
6800
Bench fee
5000
*Max. k€10 per year
†Max. k€40 voor een postdoc en k€50 voor een AiO.
Year 3
Year 4
10000
6800
2400
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
Page 11 of 11
Dossier nummer / File number: ALWNNPP2012-1-12
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 for
geese 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)
.
.1.
Novelty/originality and innovation
Judgement:
This study combines four different objectives covering the habitat use of geese, the genetic structure of a
population with a known history, the niche differentiation of three goose species breeding in the Spitsbergen and
the immunological status of breeding barnacle geese in the arctic vs. temperate regions. The combination of such
disparate approaches to study and predict future demography is particularly original, and it will lead to very
exciting results if the project team can pull all aspects together.
.
.2.
Scientific quality of the proposal
a. Scientific approach and methodology
b. Clarity and substance of objectives
c. Impact: potential extension/deepening of knowledge
d. Improvement and reinforcement of (inter)national scientific cooperation
Judgement:
Approach/Methodology
Each of the four aspects of the project seem to be sound. However, a weakness of this application is that it is not
clear how the four approaches will be brought together to achieve the ultimate aim of predicting future status of
geese in the arctic tundra.
Clarity/Substance of objectives
Each of the four objectives is clearly presented. However, how they relate together and how they will be
integrated to predict the future of geese on the arctic tundra is less clear.
Impact
Geese are key species in the arctic tundra and can have severe impacts on agricultural systems in their wintering
and migratory grounds in northern Europe. Understanding their demography and predicting future trends in goose
populations (of all species) is thus very important for future management of agricultural systems and remote arctic
regions. This project will go some way to achieving this understanding. Each of the four objectives should lead to
high quality peer-reviewed publications, and a PhD candidate will be educated in a great atmosphere between the
different aspects of the project, potentially with collaboration within the broader core program. However, the
project team will face a challenge in drawing the four objectives together to reach a prediction of the future of
geese on the tundra.
Cooperation
The project partners are from three institutions in the Netherlands with one partner from Norway, so this project
involves some but not much cooperation at the national and international level. However the research group has
a history of recent collaboration with partners in many (six) countries and if the project can build on these links in
addition it will serve to strengthen international cooperation.
.
.3.
Qualification of the applying research group
Judgement:
1
Dossier nummer / File number: ALWNNPP2012-1-12
The research group at the Arctic centre and the Netherlands research station at Ny Alesund is an internationally
recognised 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.
4.
Core programme
Each 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 are
included.
4.a.
Does the proposal fit in the core programme
Judgement:
The keystone role of geese in the arctic ecosystem, and the relationship with humans means that this project fits
very well into the core program and it seems that very close links could potentially be formed with other terrestrial
components 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 your
qualification 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 these
key species in both the arctic and the agricultural systems in Northern Europe
2
Dossier nummer / File number: ALWNNPP2012-1-12
Adviseur: 2
.
Confidential
RESEARCH PROPOSAL
(You may extend the form to have it fit your comments, a typical review has 2 or 3 pages)
.
.1.
Novelty/originality and innovation
Judgement:
Highly original and novel as it integrates key aspects of complementary disciplines in biology. Firstly, the
development of a spatially explicit goose distribution model combining data on plants and disturbance is highly
relevant for ecologist trying to anticipate the consequences of global changes on the structure and functioning of
the arctic terrestrial ecosystem. Secondly, the use of molecular techniques to study gene flow at various temporal
and spatial scale is original and relevant in the context of the actual shifts in arctic species distribution. Finally, the
project is investigating poorly known aspects of arctic species interactions (ecological niche; pathogen/parasites
effects).
.
.2.
Scientific quality of the proposal
a. Scientific approach and methodology
b. Clarity and substance of objectives
c. Impact: potential extension/deepening of knowledge
d. Improvement and reinforcement of (inter)national scientific cooperation
Judgement:
Overall assessment - HIGH QUALITY - very few research team in the world could potentially reach these original
objectives using the proposed methodology.
a. The scientific approach and methodology: convincing and novel. Considering the quality of the experts involved
in the project as well as the qualifications of the collaborators, I anticipate no major methodological issues. Note
that the project seems ambitious for a PhD student as it is highly integrative and multidisciplinary - the use of
multiple lab and modelling techniques, sources of data as well as arctic field experiment is a major strength of the
proposal 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 and
important ecological aspects in all four objectives. However, the integration of the different objectives is not always
obvious in the proposal and the addition of theoretical framework/working hypotheses for each objective would
have improved the proposal.
c. Impact - The project will certainly generate extremely relevant and new knowledge in the field of biology. I
suspect 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 will
likely contribute to the success of the proposed project. The project fit well with international research initiatives
developed recently and this will very likely strengthen the linkages established during the International Polar
Year.
.
.3.
Qualification of the applying research group
Judgement:
This is one of the best research group in the world investigating arctic wildlife ecology. The quality and quantity of
their scientific contributions are outstanding considering the challenges associated to long-term ecological
monitoring in the rapidly changing arctic ecosystem. The group perform highly relevant and innovative research
projects having strong international impacts. The team is highly interdisciplinary and can tackle challenging and
urgent research questions associated to the impact of global changes on polar regions. The team is in an
excellent position to make substantial scientific contribution in the short-term.
4.
Core programme
Each proposal forms a contribution to a core programme. The proposals are supposed to be independent,
3
Dossier nummer / File number: ALWNNPP2012-1-12
though they need to fit in the core programme. The abstracts of all proposals of the core programme are
included.
4.a.
Does the proposal fit in the core programme
Judgement:
Yes. The project will generate important interdisciplinary knowledge that strongly fit with the core program of the
Arctic Centre. The multi-faceted of the project (including immunology, field experiment, population genetic) will
place he selected PhD candidate in an excellent position to integrate knowledge produced by other team
members and to generate new data relevant to other proposals (including other proposals on arctic herbivores,
arctic plants, arctic contaminants and species interactions).
.
.
FINAL ASSESSMENT SCIENTIFIC QUALITY:
...
How do you value this proposal regarding the above mentioned qualifications? Please substantiate your
qualification in one or two concluding sentences:
- Excellent
- Very good
- Good
- Reasonable
- Poor
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 and
unique long-term data. This project will likely generate very high-quality scientific papers with substantial impact in
the international scientific community.
4
Dossier nummer / File number: ALWNNPP2012-1-12
Adviseur: 3
.
Confidential
RESEARCH PROPOSAL
(You may extend the form to have it fit your comments, a typical review has 2 or 3 pages)
.
.1.
Novelty/originality and innovation
Judgement:
I paid attention to the PhD proposal presented by Prof. M.J.J.E. Loonen on the dynamic of an arctic goose
species. This multi-disciplinary research proposal is very appealing in the sense that it would combine information
on genetic, physiology (immunology), foraging behaviour including diet, competition and habitat selection as well
as ecosystem functioning and would make use of long-term dataset and a field experiment. All these approaches
are to my point of view 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 population trends and the use by geese of the tundra.
.
.2.
Scientific quality of the proposal
a. Scientific approach and methodology
b. Clarity and substance of objectives
c. Impact: potential extension/deepening of knowledge
d. Improvement and reinforcement of (inter)national scientific cooperation
Judgement:
a.
Scientific approach and methodology
The 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 on
energetic requirements and functional responses of geese (Baveco et al. 2011). Although, this approach is very
detailed and would establish the carrying capacity for geese breeding in the Arctic, it requires a lot of parameters
that would mostly come from literature (except the functional response that will be studied in chapter 3). I question
how disturbance, predation risk, allochtonous subsidies (body reserves, lipids that contribute to egg formation in
geese, see Gauthier et al. 2003 Ecology on Snow geese) and predation (on eggs, juveniles or even adults) would
be incorporated in such models. Although, all this could probably be incorporated in such energetic modelling
framework, I would suggest the use of food-web models based on mass balance that also depend on functional
responses (i.e., DeRuiters et al. 1995 Science). It could be an alternative maybe less complicated that would have
the advantage to incorporate directly the different trophic levels of this ecosystem. This modelling approach is also
a conceptual framework (ecosystem functioning, meta ecosystems; Loreau 2011 Princeton) that would particularly
be relevant for this project and for the whole core program that attend to describe past present and future dynamic
of marine, terrestrial and freshwater ecosystems.
Objective 2. Dispersal is a key factor driving populations dynamic and still poorly understood and using molecular
techniques on an existing long-term dataset is very exciting promising. As a non-geneticist, I am not able to judge
the methodological approach presented here but the fact that preliminary analyses were already conducted is
encouraging.
Objective 3. Identifying functional responses of geese and investigate potential competition by conducting a
common garden experiment in the arctic is just lacking and it would definitely tease apart potential confounding
effects that can emerge when the niche is established on habitat selection only. I would tend to combine this
experiment with the project of Prof. Kondeur (project 06 of the core programme) and investigate stress (probably
glucocorticoids CORT on droppings I guess) of geese under the different treatment. This would give some insight
on the effect of the feeding treatment not only by measuring the body condition (based on biometric
measurements) 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 of
parasites is particularly relevant. The idea that the immune system of barnacle geese is more adaptive than the
other species is intriguing and it should deserve attention. It is not explicitly stated if such project would be
conducted in the Dutch geese populations. It would be very interesting in a perspective to explain geese
5
Dossier nummer / File number: ALWNNPP2012-1-12
migration. I also wonder if the barnacle geese that breed in the Netherlands could (partly) be issued from feral
individuals released into the wild ? This might explain their capability to mount stronger immune responses?
b. Clarity and substance of objectives
Each objectives are clear, straightforward and promising.
c. Impact: potential extension/deepening of knowledge
As stated earlier, I would tend to recommend to explore how food-web models could be another avenue to
describe trophic interactions and how perturbations (through simulations of different scenarios of resource
abundance or with the addition of other species or subsidies, see Leroux and Loreau 2008 Ecol. Lett. could modify
patterns of ecosystem functioning and in turn affect geese populations). This might be anecdotic, but the authors
should mentioned in their proposal that Svalbard terrestrial ecosystem is unique compare to other places in the
sense that there is no lemming which greatly simplify the food web and strengthen the fact that geese are key
players in the tundra. The interaction between reindeer and geese should also be of interest. Also, I found that this
project mostly emphasized on the interaction between plant and herbivores but predation can also be an important
driver in tundra ecosystems (e.g. Gilg et al. 2003 Science; Legagneux et al. 2012 Ecology).
d. Improvement and reinforcement of (inter)national scientific cooperation
This project is based on the collaboration between scientists of different horizons that would work together with
the PhD candidate. For each objective, the scientific cooperation seems relevant. I would recommend discussing
with ecosystem modellers and theoreticians that may have a different approach to address the predator-prey
interaction study. For the other objectives, the cooperation seems adequate.
.
.3.
Qualification of the applying research group
Judgement:
All the partners involved in this project are recognised international scientists. The research group is solid and
would be able to address the objectives as long as the PhD candidate is carefully selected. The PhD candidate
will indeed have to play with many concepts from various ecological disciplines (ranged from micro to macro
scales) and interact with scientists from diverse origins, which makes this project very challenging and so
interesting.
4.
Core programme
Each 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 are
included.
4.a.
Does the proposal fit in the core programme
Judgement:
.
.
FINAL ASSESSMENT SCIENTIFIC QUALITY:
...
How do you value this proposal regarding the above mentioned qualifications? Please substantiate your
qualification in one or two concluding sentences:
- Excellent
- Very good
- Good
- Reasonable
- Poor
Concluding remarks:
This proposal is ambitious and achievable because the expertise of the supervisors, mentors and their experience
of the field site is solid enough to conduct such a multidisciplinary study. For all the reasons mentioned above, I
evaluate this proposal as VERY GOOD
6
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
Title
File number
: Dr. M.J.J.E. Loonen
: From historical data to a prediction of the future for geese on arctic tundra?
: 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
1
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.
2