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Effects of cormorants on species diversity and abundance of
phantom midges and spiders by
Caroline Essenberg
Plants & Ecology Plant Ecology 2009/11 Department of Botany Stockholm University
1
Effects of cormorants on species diversity and
abundance of phantom midges and spiders
by
Caroline Essenberg
Supervisor: Peter Hambäck
Plants & Ecology
Plant Ecology 2009/11 Department of Botany Stockholm University
2
Plants & Ecology
Plant Ecology
Department of Botany
Stockholm University
S-106 91 Stockholm
Sweden
© Plant Ecology
ISSN 1651-9248
Printed by Solna Printcenter
Cover: Great cormorants (Phalacrocorax carbo) in Stockholm archipelago, Sweden.
Photo by Caroline Essenberg
3
Abstract
Great cormorant (Phalacrocorax carbo) is a colonial seabird that transports loads of nutrients
(e.g. nitrogen) from water to land. Contributed nitrogen may increase or kill the vegetation
depending on cormorant density; it may also leach into the water affecting the algal growth in
the surrounding water bodies. An increased algal growth may affect its consumers e.g.
phantom midges, which enter the terrestrial food web as predator food when swarming over
land. This study investigates how the terrestrial animal life is affected by nesting cormorants.
Phantom midges and spiders were sampled on cormorant nesting islands, abandoned
cormorant islands and non-cormorant islands to compare species richness and abundance in
two trophic levels. The study was conducted in the archipelago of Stockholm, Sweden in
2008. The statistical analysis shows a negative trend, where phantom midge species richness
decreases with higher cormorant density. Hunting and ground dwelling spider (e.g.
Lycosidae) species richness and abundance decreases significantly with higher cormorant
density. Contributed nitrogen in cormorant colonies may increase algal growth, but also cause
a shift in the primary production where fast growing plants and algae may outcompete
macrophytes. A decrease in species richness among primary producers may decrease species
richness in higher trophic levels. Spiders may also be affected negatively by changed
conditions and damaged habitats on cormorant islands and therefore decrease in species
richness and abundance.
4
Introduction
Seabirds living in both aquatic and terrestrial ecosystems are important vectors in transferring
nutrients from water to land (Lindeboom 1984; Polis and Hurd 1995, 1996; Sánches-Piñero
and Polis 2000; Osono et al. 2002; Ellis et al. 2006). When the birds feed in water, they bring
aquatic matter through their carcasses (Sanches-Pinero and Polis 2000) and guano (Anderson
and Polis 1999; Sanches-Pinero and Polis 2000; Stapp and Polis 2003; Hobara et al. 2005;
Wait et al. 2005) to their nesting places on land. The contributed nutrients are incorporated in
the soil and alter the chemical composition, indirectly affecting plants (Smith 1978; Anderson
and Polis 1999; Ellis 2005; Wait et al. 2005) and animals (Barrett et al. 2005). Seabird
populations may have important consequences for plant and animal communities on islands
and in the surrounding water bodies.
Great cormorant (Phalacrocorax carbo) is a colonial bird that is distributed almost all over
the world (Cramp and Simmons 1977). Their colonies may be very big and contain up to
10,000 breeding pairs. They catch fish in lakes, rivers and coastal areas and are one of the top-
predators in aquatic food webs (Hobson et al. 1994). By feeding in water and establishing
breeding colonies on land, cormorants transport mainly two nutrients, phosphorus and
nitrogen, from water to land (Kirkkala et al. 1998; Howarth and Marino 2006; Elser et al.
2007). Both phosphorus and nitrogen are important limiting factors of primary productivity
(Kirkkala et al. 1998; Howarth and Marino 2006; Elser et al 2007). The effects of the
increased nutrients in soil depends on cormorant nest density, it may either favour or kill
vegetation (Ellis 2005). An increased input of nitrogen and phosphorus may lead to an
increasing primary productivity (Onuf et al. 1977; Ryan & Watkins 1989; Anderson & Polis
1999; Wait et al. 2005), but when bird density is very high the nitrogen concentration in soil
becomes toxic (Ellis 2005).
Phosphorus accumulates in the surface soil, while nitrogen instead is more mobile (Vitousek
and Howarth 1991). Nitrogen in guano is uric acid and degrades microbiologically to
ammonium. Ammonium may then follow different pathways: ammonia volatilization (Hobara
et al. 2001; Lindeboom 1984), leach into the water (Wainwright et al. 1998) or be taken up by
plants (Wainwright et al. 1998; Ellis 2005). Nitrogen leaching to the water may enter the
5
marine food web through aquatic plants. It can cause an increasing growth of algae which
may affect their consumers, e.g. phantom midges (Chironomidae) (Wainwright et al. 1998).
The larval and pupal stages among phantom midges are constrained to aquatic habitats, while
the adults are aerial (Wiederholm 1989). An increase in algae growth may lead to increased
densities of phantom midges, which may re-enter the terrestrial food web as predator food.
Stable isotope analysis (a tool for studying food-web interactions) in Mellbrand (2009)
showed that spiders, which are the most abundant arthropod predators in seashores, mainly
fed on phantom midges which have fed on green algae. Hodkinson et al. (2001) found a
significant correlation between phantom midge density and spider density in coastal
ecosystems, and Sanzone et al. (2003) showed that wolf spiders (Lycosidae) heavily relied on
emerging adult insects. These studies support the hypothesis that insectivores are facilitating
the transfer of energy from emerging aquatic prey along the water edge, which in turn
increases the density of predators in seashores (Jackson and Fisher 1986).
Material and methods
Study species
Great cormorant is a species with increasing density in Europe. The number of cormorants has
increased rapidly due to eutrophication of water bodies, protective measures, reduction of
pesticides and alternation of water systems such as sluices and dams which facilitate foraging.
Overfishing has caused changes in the size distribution of fishes; large predatory fishes have
decreased which implicates an increase in the number of small fishes which enhancing the
foraging conditions of cormorants (CARSS 2002).
The increasing number of cormorants has led to conflicts between interest groups (Suter
1995). The cormorants are blamed for many different negative effects in the environment e.g.
for destruction of islands and reducing fish communities. Only a few studies investigate how
cormorants influence the terrestrial ecosystems, and the ongoing public discussion about
cormorants impact on the environment makes scientific studies about their ecological effects
very important. This study investigates how the terrestrial animal life is affected by nesting
cormorants. Abundance and species richness of phantom midges (Chironomidae) and spiders
6
(Araneae) on cormorant nesting islands and abandoned cormorant islands is analysed to
examine the effects of cormorants.
Study area
This study was conducted in the archipelago of Stockholm, Sweden. The archipelago consists
of 21,000 islands with varying sizes. The vegetation on the islands differ, some islands are
covered with bare rocks, and others with herbs, grass and even forests. Bigger islands are built
with houses, mainly summer houses.
In 2008, there were 21 cormorant colonies with 5,196 nests in the archipelago. In this study,
five cormorant nesting islands and two abandoned cormorant island were examined (table 1).
As control islands, neighbouring islands without nesting cormorants were used. Most of the
study islands were small in size and covered with grass and low vegetation. Trees were
present on most of the islands, and bigger islands contained forests. The control islands
differed in size (800-4,500 m2) from the cormorant islands, but we have tried to find
appropriate islands in relation to the study islands.
Methods
Hunting and ground dwelling spiders were captured using pitfall traps (plastic jars, 8 cm
diameter and 6 cm height) in June 2008. The jars were placed in the ground, and collected two
days after. Number of pitfall traps on each island was in relation with island size (table 1). Net
building spiders and phantom midges were sampled by sweep netting in August 2008. Sweep
netting was conducted in four gradients at two sides of the islands. Two of the gradients were
placed one meter from sea, and two gradients at ten meters from sea. Each gradient had a
distance of ten meters. Sampled spiders and phantom midges were stored in 70% alcohol until
examination at laboratory.
Adult and subadult spiders were sorted out for examination. Adult spiders were identified to
species, and subadult spiders to genus. Most of the spiders were examined by me with the
keys by Roberts (1996) and Almqvist (2005; 2006), and some spiders were sent away for
identification. Males of the sampled phantom midges were sorted into morpho-species, and
7
then sent away to Yngve Brodin at The Swedish National Environmental Protection Agency
for species identification.
On some islands, the birds pulled out some pitfall traps, which resulted in a varying number
of traps on each island (table 1) and made them therefore difficult to compare. To arrange this
problem rarefaction was calculated in the program EstimateS Win 8.2, and abundance was
calculated by dividing number of individuals with number of re-collected pitfall traps on each
island.
Linear regression was used to analyse how species richness and abundance of phantom
midges and spiders are affected by cormorant density. Cormorant density was calculated by
dividing mean number of active cormorant nests in years 2005 - 2008 with island area (m2).
All data was log-transformed and the statistical analysis was made in Statistica 5.5.
Table 1: summary of study islands.
Area Time span of Number of active cormorant nests Mean density Number of Number of
m2 colonisation 2005 2006 2007 2008 2005 - 2008 pitfall traps re-collected
(nests/m2) pitfall traps
Nesting islands
Bergskäret 22,728 1998-2008 555 595 656 637 0.0269 200 46
Småholmarna "N" 8,550 2000-2008 410 591 538 448 0.0581 100 84
Marskärskobben 7,164 2003-2008 81 186 177 212 0.0229 100 68
Ryssmasterna "N" 3,224 2003-2008 65 171 127 183 0.0423 50 24
Alörarna 2,332 2007-2008 0 0 56 156 0.0227 50 40
Abandoned islands
St. Trädskär 15,729 1996-2006 80 50 0 0 0.0021 150 75
St. Halmören 8,091 2002-2005 222 0 0 0 0.0069 100 71
Non-cormorant islands
Mjölingsö 27,285 200 190
Ägglösen 17,357 150 118
Hannas holme 7,743 100 89
Nickesören 5,209 75 34
Fårören 3,286 50 48
8
Results
Phantom midges sampled by sweep netting
In this study, 6,141 male phantom midges were caught. A varying number of individuals were
caught on the different island categories. On cormorant nesting islands, 4,942 individuals
were caught, 140 individuals on abandoned cormorant islands and 1,059 individuals on non-
cormorant islands (app. 1, table 1). The big difference in the number of sampled phantom
midges is caused by the high number of individuals caught on the cormorant nesting island,
Alörarna, where 4,039 individuals were caught.
Species determination resulted in 41 different species (app. 1, table 1). The number of species
on each island varied between 9 and 24 species, and the most abundant species on the study
islands were Paratanytarsus dissimilis, Tanytarsus usmaensis, Cricotopus bicinctus and
Cricotopus caducus. Most of the sampled species are common in Sweden and Europe except
for C. caducus, which was found in Sweden for the first time. The species is common in the
rest of Europe and in my samples it was one of the four most abundant species. It was found
on most of the study islands and is probably a quite common species in the rest of the
archipelago. Linear regression analysis of cormorant density and phantom midges species
richness (R2=0.32, F=4.78, N=12, p=0.054) (fig. 1), and abundance and cormorant density
(R2=0.12, F=1.33, N=12, p=0.280) were not significant.
9
Cormorant density (nests/m2)
Num
ber
of
specie
s
(scale
is log-t
ransfo
rmed)
2.0
2.2
2.4
2.6
2.8
3.0
3.2
3.4
0.00 0.01 0.02 0.03 0.04 0.05 0.06
Figure 1: Species richness of phantom midges.
Spiders sampled by pitfall traps
In a total, 39 spider species (app. 1, table 2) and 598 individuals were sampled. The most
abundant species on the study islands were Pardosa amentata, Pardosa agricola, Trochosa
ruricola and Pachygnatha degeeri. On most of the islands, P. amentata was the most
abundant species, but P. agricola was very common on three of the study islands. Many
spiders of the group “money spiders” were also caught in the pitfall traps. Because of the
difficulty of species identification of these spiders, they were all sorted out and are not
included in this study.
On cormorant nesting islands 92 individuals were caught, 206 individuals on abandoned
cormorant islands and 300 individuals on non-cormorant islands (app.1, table 2). Rarefaction
resulted in a variation between 1 and 9.53 species on each island. Linear regression analysis
shows that abundance (R2=0.47, F=8.76, N=12, p=0.014) (fig. 2) and species richness
(R2=0.56, F=12.67, N=12, p=0.005) (fig. 3) of hunting and ground dwelling spiders decreases
with higher cormorant density.
Two points
10
Cormorant density (nests/m2)
Ab
un
da
nce
(sca
le is lo
g-t
ran
sfo
rme
d)
-3,5
-3,0
-2,5
-2,0
-1,5
-1,0
-0,5
0,0
0,5
1,0
0,00 0,01 0,02 0,03 0,04 0,05 0,06
Figure 2: Abundance of hunting and ground dwelling spiders (e.g. Lycosidae). Abundance
was estimated on number of individuals / pitfall trap.
Cormorant density (nests/m2)
Nu
mb
er
of
sp
ecie
s
(sca
le is lo
g-t
ran
sfo
rme
d)
0,0
0,4
0,8
1,2
1,6
2,0
2,4
0,00 0,01 0,02 0,03 0,04 0,05 0,06
Figure 3: Species richness of hunting and ground dwelling spiders (e.g. Lycosidae). Number
of species is rarefied.
Two points
11
Spiders sampled by sweep netting
With sweep netting, 344 individuals and 21 species were sampled. Species list can be found in
appendix 1, table 3. Tetragnatha montana was the most abundant species on the study islands,
Larinioides cornutus, Tetragnatha dearmata and Xysticus cristatus were also common
species. On cormorant nesting islands 119 individuals were caught, 85 individuals on
abandoned cormorant islands and 140 individuals were caught on non-cormorant islands (app.
1, table 3). Linear regression analysis of species richness and cormorant density (R2=0.03,
F=0.27, N=12, p=0.616), and abundance and cormorant density (R2=0.009, F=0.09, N=12,
p=0.771) were not significant.
Correlation analysis
To test if hunting and ground dwelling spider species richness are depending on phantom
midge species richness, a correlation analysis was made (r=0.57, p=0.053) (fig. 4).
(scale is log-transformed)
Number of spiders species
Nu
mb
er
of
ph
an
tom
mid
ge
sp
ecie
s
(sca
le is lo
g-t
ran
sfo
rme
d)
2,0
2,2
2,4
2,6
2,8
3,0
3,2
3,4
0,0 0,4 0,8 1,2 1,6 2,0 2,4
Figure 4: Correlation analysis of phantom midges species richness and hunting and ground
dwelling spider (e.g. Lycosidae) species richness. Spider species richness is rarefied.
12
Discussion
The cormorants bring loads of nutrients to their nesting places, affecting the surrounding
environment. They nest in big colonies on small areas and bring nutrients (phosphorus and
nitrogen) from water to land (Kirkkala et al. 1998; Howarth and Marino 2006; Elser et al.
2007). The nutrients incorporate in soil, and nitrogen may also leach into the water and affect
the aquatic plants and indirectly its consumers (Wainright et al. 1998). Contributed nitrogen
may increase algal growth (Wainright et al. 1998), but also cause a shift in primary producers.
Fast growing macroalgae, epiphytes and phytoplankton will out-compete macrophytes by
exploiting the nutrients and by reducing light penetration for submerged plants (Sand-Jensen
and Borum 1991; Short et al. 1995; Taylor et al. 1995; Wear et al. 1999). High species
diversity among primary producers allows rare consumer species to become abundant and,
therefore, increases species richness of the consumers (Hutchinson 1959). A decrease in
diversity of primary production consequently leads to a decrease in consumer diversity
(Siemann 1998; Haddad et al. 2000). This study shows a negative trend (p=0,054) between
phantom midge species richness and cormorant density probably caused by a decrease in
diversity of primary producers due to a high input of nitrogen.
Phantom midges are aerial as adults (Wiederholm 1989) and enter the terrestrial system when
swarming over land (Wainright et al. 1998). Spiders are the most abundant arthropod
predators on seashores and mainly feed on phantom midges (Jackson and Fisher 1986;
Hodkinson et al. 2001; Sanzone et al. 2003; Mellbrand 2009). A decrease in herbivore
diversity may also decrease the diversity in higher trophic levels (Siemann 1998; Keats et al.
2004), and this study shows that hunting and ground dwelling spiders (e.g. Lycosidae)
significantly decreases in abundance and species richness with higher cormorant density.
Another important factor that may affect species diversity and abundance of spiders on
cormorant islands is the terrestrial vegetation. When the high input of nutrients is incorporated
in the soil, it affects the terrestrial plants by increasing biomass and by decreasing species
richness (Smith 1978; Anderson and Polis 1999; Ellis 2005; Wait et al. 2005). In seabird
colonies, annual and biennial plant species increase while perennial species decrease (Smith
1978; Vidal et a. 1998; Garcia et al. 2002), and when the bird densities are very high the
nutrient concentration in soil becomes toxic and may kill the vegetation (Ellis 2005). Spiders
13
may be sensitive to changed conditions and damage habitats and consequently decrease in
abundance and species richness (Greenstone 1984; Siemann 1998; Haddad et al. 2000).
Net building spiders showed no significant response to cormorant density in abundance and
species richness. This is quite unexpected since these spiders may as well as hunting spiders
utilize phantom midges as food resource to a high degree and persists in reduced vegetation.
Web building spiders differ in hunting methods and ecology from ground dwelling spiders
and may respond differently to damage habitats.
Conclusion
The cormorants bring loads of nutrients to their nesting places which affect the environment
around them. Cormorant derived nitrogen leaching to the water affects the surrounding water
bodies by changing species composition of aquatic plants. This study indicates that species
richness of phantom midges (algae consumers) decreases with higher cormorant density as a
result of low species richness among alage around cormorant islands. Phantom midges are
aerial as adults and enter the terrestrial food web as predator food. Hunting and ground
dwelling spiders (e.g. Lycosidae), which are common arthropod predators in seashores,
decreases significantly in abundance and species richness with higher cormorant density. This
may be a result of a decreased species diversity of prey and damaged habitats on cormorant
islands.
Acknowledgements
I want to thank Peter Hambäck and Gundula Kolb for supervising this study, and Lenn and
Eskil Jerling for their hospitality and helping in the field. I also want to thank Yngve Brodin
for species identification of phantom midges, and Sandra Öberg, Torbjörn Kronstedt and Lars
Jonsson for helping with species identification of spiders.
14
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17
Appendix 1
Table 1: list of phantom midge species and number of individuals
Question mark between genus and species signifies that the genus is assured but not species.
Subfamily Species Cormorant Abandoned Non - Total
nesting cormorant cormorant
islands islands island
Chironominae Chironomus aprilinus 228 22 222 472
Chironomus plumosus 12 5 10 27
Dicrotendipes pulsus 213 4 78 295
Harnischia curtilamellata 1 1
Microchironomus tener 13 14 27
Microtendipes pedellus 220 26 57 303
Parachironomus arcuatus 7 8 15
Paratanytarsus dissimilis 466 10 139 615
Paratanytarsus natvigi 3 3
Paratendipes albimanus 2 2
Phaenopsectra flavipes 20 25 45
Polypedilum nubeculosum 52 36 88
Polypedilum scalaenum 253 40 24 317
Tanytarsus ? brundini 134 40 174
Tanytarsus usmaensis 927 4 74 1005
Xenochironomus xenolabis 11 4 15
Orthocladiinae Bryophaenocladius ? scanicus 5 11 16
Cricotopus annulator 4 4
Cricotopus bicinctus 1618 2 92 1712
Cricotopus caducus 458 58 516
Cricotopus flavocinctus 1 1
Cricotopus intersectus 10 8 18
Cricotopus ? ornatus 2 2
Cricotopus sylvestris 119 2 63 184
Halocladius variabilis 30 4 2 36
Limnophyes difficilis 6 23 29
Limnophyes minimus 3 3
Limnophyes pumilio 3 6 9
Limnophyes sp. A 1 1
Nanocladius dichromus 4 4
Nanocladius rectinervis 1 1
Orthocladius sp. A 1 1
Paratrichocladius rufiventris 6 1 7
Psectrocladius ? limbatellus 1 1
Psectrocladius ? oligosetus 7 7
Smittia sp. A 1 1
Tanypodiane Ablabesmyia monilis 56 13 69
Ablabesmyia longistyla 2 2
18
Procladius culiciformis 10 14 16 40
Procladius ? ferrugineus 4 4 7 15
Procladius ? sagittalis 41 3 14 58
Table 2: list of spiders species and number of individuals sampled by pitfall traps
Family Species Cormorant Abandoned Non - Total
nesting cormorant cormorant
islands islands island
Araneidae Larinioides patagiatus 1 1 2
Larinioides (imm.) 2 2
Clubionidae Clubiona (imm.) 1 1 2
Gnaphosidae Callilepis nocturna 3 3
Drassyllus praeficus 1 1
Gnaphosa montana 1 1
Haplodrassus signifer 3 3
Micaria nimosa 1 1
Micaria pulicaria 2 1 3
Zelotes petrensis 1 1
Zelotes subterraneus 2 1 5 8
Zelotes (imm.) 4 1 4 9
Hahniidae Antistea elegans 1 1
Linyphiidae Lepthyphantes flavipes 1 1 2
Lepthyphantes mengei 1 1
Stemonyphantes lineatus 1 1
Tenuiphantes flavipes 1 1
Lycosidae Alopecosa pulverulenta 8 8
Alopecosa (imm.) 1 1
Arctosa leopardus 1 1 10 12
Arctosa (imm.) 6 7 13
Pardosa agrestis 2 1 7 10
Pardosa agricola 14 49 80 143
Pardosa amentata 8 34 17 59
Pardosa fulvipes 3 5 8
Pardosa lugubris 3 3
Pardosa monticola 1 12 13
Pardosa nigriceps 2 2
Pardosa palustris 2 1 3
Pardosa prativaga 12 17 29
Pardosa pullata 2 2 4
Pardosa (imm.) 3 3
Pirata piraticus 1 1
19
Trochosa ruricola 9 10 10 29
Trochosa spinipalpis 1 1 2
Trochosa terricola 2 4 4 10
Trochosa (imm.) 8 31 33 72
Salticidae Heliophanus cupreus 1 1
Tetragnathidae Pachygnatha clercki 2 17 2 21
Pachygnatha degeeri 7 49 45 101
Theridiidae Crustulina guttata 1 1
Thomisidae Ozyptila praticola 3 3
Thanatus striatus 1 1
Xysticus cristatus 2 2
Zoridae Zora spinimana 1 1
Table 3: list of spider species and number of individuals sampled by sweep netting
Family Species Cormorant Abandoned Non - Total
nesting cormorant cormorant
islands islands island
Araneidae Araniella (imm.) 2 2
Larinioides cornutus 2 4 1 7
Larinioides patagiatus 1 1 2
Larinioides (imm.) 20 21 9 50
Clubionidae Clubiona (imm.) 3 3
Dictynidae Dictyna (imm.) 7 7
Linyphiidae Gongyliduim rufipes 3 3
Kaestneria dorsalis 5 8 13
Neriene (imm.) 1 1
Salticidae Heliophanus dubius 1 1 2
Heliophanus (imm.) 5 5
Tetragnathidae Meta mengei 1 1
Meta segmentata 9 1 10 20
Meta (imm.) 1 1 2
Pachygnatha degeeri 1 1
Tetragnatha dearmata 12 6 5 23
Tetragnatha extensa 2 4 6 12
Tetragnatha montana 32 24 18 74
Tetragnatha striata 3 1 11 15
Tetragnatha (imm.) 16 5 29 50
Theridiidae Anelosimus (imm.) 1 1
Theridion (imm.) 1 2 3
Thomisidae Philodromus (imm.) 2 4 6
Tibellus oblongus 1 1
20
Xysticus cristatus 1 1 4 6
Xysticus (imm.) 5 16 13 34
21
Serien Plants & Ecology (ISSN 1651-9248) har tidigare haft namnen "Meddelanden från
Växtekologiska avdelningen, Botaniska institutionen, Stockholms Universitet" nummer
1978:1 – 1993:1 samt "Växtekologi". (ISSN 1400-9501) nummer 1994:1 – 2003:3.
Följande publikationer ingår i utgivningen:
1978:1 Liljelund, Lars-Erik: Kompendium i matematik för ekologer.
1978:2 Carlsson, Lars: Vegetationen på Littejåkkadeltat vid Sitasjaure, Lule Lappmark.
1978:3 Tapper, Per-Göran: Den maritima lövskogen i Stockholms skärgård.
1978:4: Forsse, Erik: Vegetationskartans användbarhet vid detaljplanering av
fritidsbebyggelse.
1978:5 Bråvander, Lars-Gunnar och Engelmark, Thorbjörn: Botaniska studier vid
Porjusselets och St. Lulevattens stränder i samband med regleringen 1974.
1979:1 Engström, Peter: Tillväxt, sulfatupptag och omsättning av cellmaterial hos
pelagiska saltvattensbakterier.
1979:2 Eriksson, Sonja: Vegetationsutvecklingen i Husby-Långhundra de senaste
tvåhundra åren.
1979:3 Bråvander, Lars-Gunnar: Vegetation och flora i övre Teusadalen och vid Auta-
och Sitjasjaure; Norra Lule Lappmark. En översiktlig inventering med anledning av
områdets exploatering för vattenkraftsändamål i Ritsemprojektet.
1979:4 Liljelund, Lars-Erik, Emanuelsson, Urban, Florgård, C. och Hofman-Bang,
Vilhelm: Kunskapsöversikt och forskningsbehov rörande mekanisk påverkan på
mark och vegetation.
1979:5 Reinhard, Ylva: Avloppsinfiltration - ett försök till konsekvensbeskrivning.
1980:1 Telenius, Anders och Torstensson, Peter: Populationsstudie på Spergularia marina
och Spergularia media. I Frödimorfism och reproduktion.
1980:2 Hilding, Tuija: Populationsstudier på Spergularia marina och Spergularia media.
II Resursallokering och mortalitet.
1980:3 Eriksson, Ove: Reproduktion och vegetativ spridning hos Potentilla anserina L.
1981:1 Eriksson, Torsten: Aspekter på färgvariation hos Dactylorhiza sambucina.
1983:1 Blom, Göran: Undersökningar av lertäkter i Färentuna, Ekerö kommun.
1984:1 Jerling, Ingemar: Kalkning som motåtgärd till försurningen och dess effekter på
blåbär, Vaccinium myrtillus.
1986:1 Svanberg, Kerstin: En studie av grusbräckans (Saxifraga tridactylites) demografi.
1986:2 Nyberg, Hans: Förändringar i träd- och buskskiktets sammansättning i
ädellövskogen på Tullgarnsnäset 1960-1983.
1987:1 Edenholm, Krister: Undersökningar av vegetationspåverkan av vildsvinsbök i
Tullgarnsområdet.
1987:2 Nilsson, Thomas: Variation i fröstorlek och tillväxthastighet inom släktet Veronica.
1988:1 Ehrlén, Johan: Fröproduktion hos vårärt (Lathyrus vernus L.). - Begränsningar och
reglering.
1988:2 Dinnétz, Patrik: Local variation in degree of gynodioecy and protogyny in Plantago
maritima.
1988:3 Blom, Göran och Wincent, Helena: Effekter of kalkning på ängsvegetation.
1989:1 Eriksson, Pia: Täthetsreglering i Littoralvegetation.
1989:2 Kalvas, Arja: Jämförande studier av Fucus-populationer från Östersjön och
västkusten.
1990:1 Kiviniemi, Katariina: Groddplantsetablering och spridning hos smultron, Fragaria
vesca.
22
1990:2 Idestam-Almquist, Jerker: Transplantationsförsök med Borstnate.
1992:1 Malm, Torleif: Allokemisk påverkan från mucus hos åtta bruna makroalger på
epifytiska alger.
1992:2 Pontis, Cristina: Om groddknoppar och tandrötter. Funderingar kring en klonal
växt: Dentaria bulbifera.
1992:3 Agartz, Susanne: Optimal utkorsning hos Primula farinosa.
1992:4 Berglund, Anita: Ekologiska effekter av en parasitsvamp - Uromyces lineolatus på
Glaux maritima (Strandkrypa).
1992:5 Ehn, Maria: Distribution and tetrasporophytes in populations of Chondrus crispus
Stackhouse (Gigartinaceae, Rhodophyta) on the west coast of Sweden.
1992:6 Peterson, Torbjörn: Mollusc herbivory.
1993:1 Klásterská-Hedenberg, Martina: The influence of pH, N:P ratio and zooplankton
on the phytoplanctic composition in hypertrophic ponds in the Trebon-region, Czech
Republic.
1994:1 Fröborg, Heléne: Pollination and seed set in Vaccinium and Andromeda.
1994:2 Eriksson, Åsa: Makrofossilanalys av förekomst och populationsdynamik hos Najas
flexilis i Sörmland.
1994:3 Klee, Irene: Effekter av kvävetillförsel på 6 vanliga arter i gran- och tallskog.
1995:1 Holm, Martin: Beståndshistorik - vad 492 träd på Fagerön i Uppland kan berätta.
1995:2 Löfgren, Anders: Distribution patterns and population structure of an economically
important Amazon palm, Jessenia bataua (Mart.) Burret ssp. bataua in Bolivia.
1995:3 Norberg, Ylva: Morphological variation in the reduced, free floating Fucus
vesiculosus, in the Baltic Proper.
1995:4 Hylander, Kristoffer & Hylander, Eva: Mount Zuquala - an upland forest of
Ethiopia. Floristic inventory and analysis of the state of conservation.
1996:1 Eriksson, Åsa: Plant species composition and diversity in semi-natural grasslands -
with special emphasis on effects of mycorrhiza.
1996:2 Kalvas, Arja: Morphological variation and reproduction in Fucus vesiculosus L.
populations.
1996:3 Andersson, Regina: Fågelspridda frukter kemiska och morfologiska egenskaper i
relation till fåglarnas val av frukter.
1996:4 Lindgren, Åsa: Restpopulationer, nykolonisation och diversitet hos växter i
naturbetesmarker i sörmländsk skogsbygd.
1996:5 Kiviniemi, Katariina: The ecological and evolutionary significance of the early life
cycle stages in plants, with special emphasis on seed dispersal.
1996:7 Franzén, Daniel: Fältskiktsförändringar i ädellövskog på Fagerön, Uppland,
beroende på igenväxning av gran och skogsavverkning.
1997:1 Wicksell, Maria: Flowering synchronization in the Ericaceae and the Empetraceae.
1997:2 Bolmgren, Kjell: A study of asynchrony in phenology - with a little help from
Frangula alnus.
1997:3 Kiviniemi, Katariina: A study of seed dispersal and recruitment of plants in a
fragmented habitat.
1997:4 Jakobsson, Anna: Fecundity and abundance - a comparative study of grassland
species.
1997:5 Löfgren, Per: Population dynamics and the influence of disturbance in the Carline
Thistle, Carlina vulgaris.
1998:1 Mattsson, Birgitta: The stress concept, exemplified by low salinity and other stress
factors in aquatic systems.
23
1998:2 Forsslund, Annika & Koffman, Anna: Species diversity of lichens on decaying
wood - A comparison between old-growth and managed forest.
1998:3 Eriksson, Åsa: Recruitment processes, site history and abundance patterns of plants
in semi-natural grasslands.
1998:4 Fröborg, Heléne: Biotic interactions in the recruitment phase of forest field layer
plants.
1998:5 Löfgren, Anders: Spatial and temporal structure of genetic variation in plants.
1998:6 Holmén Bränn, Kristina: Limitations of recruitment in Trifolium repens.
1999:1 Mattsson, Birgitta: Salinity effects on different life cycle stages in Baltic and North
Sea Fucus vesiculosus L.
1999:2 Johannessen, Åse: Factors influencing vascular epiphyte composition in a lower
montane rain forest in Ecuador. An inventory with aspects of altitudinal distribution,
moisture, dispersal and pollination.
1999:3 Fröborg, Heléne: Seedling recruitment in forest field layer plants: seed production,
herbivory and local species dynamics.
1999:4 Franzén, Daniel: Processes determining plant species richness at different scales -
examplified by grassland studies.
1999:5 Malm, Torleif: Factors regulating distribution patterns of fucoid seaweeds. A
comparison between marine tidal and brackish atidal environments.
1999:6 Iversen, Therese: Flowering dynamics of the tropical tree Jacquinia nervosa.
1999:7 Isæus, Martin: Structuring factors for Fucus vesiculosus L. in Stockholm south
archipelago - a GIS application.
1999:8 Lannek, Joakim: Förändringar i vegetation och flora på öar i Norrtälje skärgård.
2000:1 Jakobsson, Anna: Explaining differences in geographic range size, with focus on
dispersal and speciation.
2000:2 Jakobsson, Anna: Comparative studies of colonisation ability and abundance in
semi-natural grassland and deciduous forest.
2000:3 Franzén, Daniel: Aspects of pattern, process and function of species richness in
Swedish seminatural grasslands.
2000:4 Öster, Mathias: The effects of habitat fragmentation on reproduction and population
structure in Ranunculus bulbosus.
2001:1 Lindborg, Regina: Projecting extinction risks in plants in a conservation context.
2001:2 Lindgren, Åsa: Herbivory effects at different levels of plant organisation; the
individual and the community.
2001:3 Lindborg, Regina: Forecasting the fate of plant species exposed to land use change.
2001:4 Bertilsson, Maria: Effects of habitat fragmentation on fitness components.
2001:5 Ryberg, Britta: Sustainability aspects on Oleoresin extraction from Dipterocarpus
alatus.
2001:6 Dahlgren, Stefan: Undersökning av fem havsvikar i Bergkvara skärgård, östra
egentliga Östersjön.
2001:7 Moen, Jon; Angerbjörn, Anders; Dinnetz, Patrik & Eriksson Ove: Biodiversitet i
fjällen ovan trädgränsen: Bakgrund och kunskapsläge.
2001:8 Vanhoenacker, Didrik: To be short or long. Floral and inflorescence traits of Bird`s
eye primrose Primula farinose, and interactions with pollinators and a seed predator.
2001:9 Wikström, Sofia: Plant invasions: are they possible to predict?
2001:10 von Zeipel, Hugo: Metapopulations and plant fitness in a titrophic system – seed
predation and population structure in Actaea spicata L. vary with population size.
2001:11 Forsén, Britt: Survival of Hordelymus europaéus and Bromus benekenii in a
deciduous forest under influence of forest management.
24
2001:12 Hedin, Elisabeth: Bedömningsgrunder för restaurering av lövängsrester i Norrtälje
kommun.
2002:1 Dahlgren, Stefan & Kautsky, Lena: Distribution and recent changes in benthic
macrovegetation in the Baltic Sea basins. – A literature review.
2002:2 Wikström, Sofia: Invasion history of Fucus evanescens C. Ag. in the Baltic Sea
region and effects on the native biota.
2002:3 Janson, Emma: The effect of fragment size and isolation on the abundance of Viola
tricolor in semi-natural grasslands.
2002:4 Bertilsson, Maria: Population persistance and individual fitness in Vicia pisiformis:
the effects of habitat quality, population size and isolation.
2002:5 Hedman, Irja: Hävdhistorik och artrikedom av kärlväxter i ängs- och hagmarker på
Singö, Fogdö och norra Väddö.
2002:6 Karlsson, Ann: Analys av florans förändring under de senaste hundra åren, ett
successionsförlopp i Norrtälje kommuns skärgård.
2002:7 Isæus, Martin: Factors affecting the large and small scale distribution of fucoids in
the Baltic Sea.
2003:1 Anagrius, Malin: Plant distribution patterns in an urban environment, Södermalm,
Stockholm.
2003:2 Persson, Christin: Artantal och abundans av lavar på askstammar – jämförelse
mellan betade och igenvuxna lövängsrester.
2003:3 Isæus, Martin: Wave impact on macroalgal communities.
2003:4 Jansson-Ask, Kristina: Betydelsen av pollen, resurser och ljustillgång för
reproduktiv framgång hos Storrams, Polygonatum multiflorum.
2003:5 Sundblad, Göran: Using GIS to simulate and examine effects of wave exposure on
submerged macrophyte vegetation.
2004:1 Strindell, Magnus: Abundansförändringar hos kärlväxter i ädellövskog – en
jämförelse av skötselåtgärder.
2004:2 Dahlgren, Johan P: Are metapopulation dynamics important for aquatic plants?
2004:3 Wahlstrand, Anna: Predicting the occurrence of Zostera marina in bays in the
Stockholm archipelago,northern Baltic proper.
2004:4 Råberg, Sonja: Competition from filamentous algae on Fucus vesiculosus –
negative effects and the implications on biodiversity of associated flora and fauna.
2004:5 Smaaland, John: Effects of phosphorous load by water run-off on submersed plant
communities in shallow bays in the Stockholm archipelago.
2004:6 Ramula Satu: Covariation among life history traits: implications for plant
population dynamics.
2004:7 Ramula, Satu: Population viability analysis for plants: Optimizing work effort and
the precision of estimates.
2004:8 Niklasson, Camilla: Effects of nutrient content and polybrominated phenols on the
reproduction of Idotea baltica and Gammarus ssp.
2004:9 Lönnberg, Karin: Flowering phenology and distribution in fleshy fruited plants.
2004:10 Almlöf, Anette: Miljöfaktorers inverkan på bladmossor i Fagersjöskogen, Farsta,
Stockholm.
2005:1 Hult, Anna: Factors affecting plant species composition on shores - A study made in
the Stockholm archipelago, Sweden.
2005:2 Vanhoenacker, Didrik: The evolutionary pollination ecology of Primula farinosa.
2005:3 von Zeipel, Hugo: The plant-animal interactions of Actea spicata in relation to
spatial context.
2005:4 Arvanitis, Leena T.: Butterfly seed predation.
25
2005:5 Öster, Mathias: Landscape effects on plant species diversity – a case study of
Antennaria dioica.
2005:6 Boalt, Elin: Ecosystem effects of large grazing herbivores: the role of nitrogen.
2005:7 Ohlson, Helena: The influence of landscape history, connectivity and area on
species diversity in semi-natural grasslands.
2005:8 Schmalholz, Martin: Patterns of variation in abundance and fecundity in the
endangered grassland annual Euphrasia rostkovia ssp. Fennica.
2005:9 Knutsson, Linda: Do ants select for larger seeds in Melampyrum nemorosum?
2006:1 Forslund, Helena: A comparison of resistance to herbivory between one exotic and
one native population of the brown alga Fucus evanescens.
2006:2 Nordqvist, Johanna: Effects of Ceratophyllum demersum L. on lake phytoplankton
composition.
2006:3 Lönnberg, Karin: Recruitment patterns, community assembly, and the evolution of
seed size.
2006:4 Mellbrand, Kajsa: Food webs across the waterline - Effects of marine subsidies on
coastal predators and ecosystems.
2006:5 Enskog, Maria: Effects of eutrophication and marine subsidies on terrestrial
invertebrates and plants.
2006:6 Dahlgren, Johan: Responses of forest herbs to the environment.
2006:7 Aggemyr, Elsa: The influence of landscape, field size and shape on plant species
diversity in grazed former arable fields.
2006:8 Hedlund, Kristina: Flodkräftor (Astacus astacus) i Bornsjön, en omnivors påverkan
på växter och snäckor.
2007:1 Eriksson, Ove: Naturbetesmarkernas växter- ekologi, artrikedom och
bevarandebiologi.
2007:2 Schmalholz, Martin: The occurrence and ecological role of refugia at different
spatial scales in a dynamic world.
2007:3 Vikström, Lina: Effects of local and regional variables on the flora in the former
semi-natural grasslands on Wäsby Golf club’s course.
2007:4 Hansen, Joakim: The role of submersed angiosperms and charophytes for aquatic
fauna communities.
2007:5 Johansson, Lena: Population dynamics of Gentianella campestris, effects of
grassland management, soil conditions and the history of the landscape
2007:6 von Euler, Tove: Sex related colour polymorphism in Antennaria dioica.
2007:7 Mellbrand, Kajsa: Bechcombers, landlubbers and able seemen: Effects of marine
subsidies on the roles of arthropod predators in coastal food webs.
2007:8 Hansen, Joakim: Distribution patterns of macroinvertebrates in vegetated, shallow,
soft-bottom bays of the Baltic Sea.
2007:9 Axemar, Hanna: An experimental study of plant habitat choices by
macroinvertebrates in brackish soft-bottom bays.
2007:10 Johnson, Samuel: The response of bryophytes to wildfire- to what extent do they
survive in-situ?
2007:11 Kolb, Gundula: The effects of cormorants on population dynamics and food web
structure on their nesting islands.
2007:12 Honkakangas, Jessica: Spring succession on shallow rocky shores in northern
Baltic proper.
2008:1 Gunnarsson, Karl: Påverkas Fucus radicans utbredning av Idotea baltica?
2008:2 Fjäder, Mathilda: Anlagda våtmarker i odlingslandskap- Hur påverkas
kärlväxternas diversitet?
26
2008:3 Schmalholz, Martin: Succession in boreal bryophyte communities – the role of
microtopography and post-harvest bottlenecks.
2008:4 Jokinen, Kirsi: Recolonization patterns of boreal forest vegetation following a
severe flash flood.
2008:5 Sagerman, Josefin: Effects of macrophyte morphology on the invertebrate fauna in
the Baltic Sea.
2009:1 Andersson, Petter: Quantitative aspects of plant-insect interaction in fragmented
landscapes – the role of insect search behaviour.
2009:2 Kolb, Gundula: The effects of cormorants on the plant-arthropod food web on their
nesting islands.
2009:3 Johansson, Veronika: Functional traits and remnant populations in abandoned
semi-natural grasslands.
2009: 4 König, Malin: Phenotypic selection on flowering phenology and herbivory in
Cardamine amara.
2009:5 Forslund, Helena: Grazing and the geographical range of seaweeds –
The introduced Fucus evanescens and the newly described Fucus radicans.
2009:6 von Euler, Tove: Local adaptation and life history differentiation in plant
populations.
2009:7 Tiderman, Dan: Sympatric speciation in Baltic Sea Fucus population- Is vegetative
reproduction the key for evolution of F.radicans?
2009:8 Marteinsdóttir, Bryndis: Asembly of plant communties in grasslands: an
overview.
2009:9 Herrström, Anna: Distribution and host plant selection of the Alcon blue
(Maculinea alcon).
2009:10 Norrman, Karolina: Long-term exclusion of mammalian herbivores affect plant
biomass in plant communities of the tundra in northern Norway.
27