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Supplementary Online Material:
Global change and species interactions in terrestrial
ecosystems
Jason M. Tylianakis, Raphael K. Didham, Jordi Bascompte, David A. Wardle
Contents:
Box S1: Higher order effects of multiple GEC drivers on species interactions.
Box S2: Effects of Global Environmental Change (GEC) drivers on biotic interactions.
Box S3: The consequences of interaction network architecture for the effects of GEC on
two different species interaction types.
Methods S1: Description of methods and search terms used for literature review.
Figures S1-S6: Additional information supporting the trends in changes to pairwise
interactions depicted in main text Fig. 1.
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Box S1: Higher order effects of multiple GEC drivers on species interactions.
As an illustrative example of the potential higher order interactions among multiple drivers of
GEC, we show the relative effects of three GEC drivers (CO2 enrichment, climate change
and nitrogen deposition) on one common type of species interaction (foliar herbivory).
Arrows a, b and c are direct effects, whereas arrows d, e, and f moderate the direct effects (of
a, b and c respectively).
Climate change may have direct effects on herbivory (arrow b), with performance
(Johns & Hughes 2002; Zvereva & Kozlov 2006) and abundance (Bezemer et al. 1998;
Logan et al. 2003) of herbivores tending to improve under conditions of simulated climate
change, and ranges of herbivore species potentially expanding (Battisti et al. 2005). In
contrast, the direct effects of elevated CO2 on herbivory (arrow c) may often be negative,
with herbivore performance and fecundity often decreasing under conditions of elevated CO2
(Percy et al. 2002; Awmack et al. 2004; Asshoff & Hättenschwiler 2005; Zvereva & Kozlov
2006). Carbon dioxide enrichment may also drive climate change (arrow a), such that the
effect of CO2 is the sum of the direct (c) and climate-mediated indirect (in a path model,
coefficient a x b) effects on herbivory. Furthermore, the effects of climate change and CO2
enrichment may be moderated by N deposition (dotted arrows). For example, reduced
herbivory of some plant species under elevated CO2 may be compensated for by increasing
herbivory under N fertilisation (arrow f), potentially producing a positive interaction effect
between these two drivers when they are tested simultaneously (Cleland et al. 2006). Plant
growth under elevated CO2 becomes increasingly N limited, so N deposition may increase
plant biomass production and thereby reduce the impact of CO2 on climate change (arrow d).
However, these short-term benefits to plant growth will themselves be largely offset by
climate change (Long et al. 2006). Finally, N deposition and temperature increases usually
have positive effects on herbivory, and these effects can be additive (Richardson et al. 2002)
or interactive (e) (Ritchie 2000).
N deposition
d
e
f
Climate
b
a
CO2
c
Plant-Herbivore
Interaction
Above: Potential higher order interactions among CO2 enrichment, climate change and
nitrogen deposition effects on herbivory. Direct effects of N deposition on herbivory are
omitted, but examples of these are presented in Fig. S2.
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Box S2: Effects of Global Environmental Change (GEC) drivers on biotic
interactions. GEC drivers may directly affect (block red arrows) the phenotype (e.g.
physiology) or abundance of an organism that indirectly affects (black arrows) the
organism’s interactions with other species (its consumers or resources). For taxa on
which the direct effects of GEC are known to be trivial or unimportant (e.g., direct
CO2 effects on organisms other than plants) interactions are not depicted. GEC drivers
may also directly modify the interaction between two organisms (red, dotted arrows)
e.g. through changes in phenology of both interacting species. Double headed arrows
indicate competitive interactions. For simplicity, competitive interactions are only
depicted between plants, and between native and exotic species, but species at any
trophic level may compete with others.
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Box S3: The consequences of interaction network architecture for the effects of GEC on
two different species interaction types.
Mutualistic networks are very cohesive and show a nested structure in which
specialist species interactions form well-defined subsets of the interactions between
generalist species (bottom left). These features make mutualist networks robust to the
loss of interactions caused by GEC (the specialist animals that pollinate fewest plants
pollinate a subset of the plants pollinated by generalists). Antagonistic networks
appear to be more compartmentalised, with each compartment including a group of
strongly interacting plants and animals, but few interactions among different
compartments (bottom right) (Lewinsohn et al. 2006). This structure makes them
more susceptible to losses of species, especially top consumers. However, recent work
on ant-plant mutualist networks has shown that symbiotic associations (e.g. ants and
myrmecophytes) are highly compartmentalised and less nested than nonsymbiotic
(e.g. seed dispersal) networks (Guimarães et al. 2007), indicating that tightly
coevolved symbioses may be particularly prone to extinctions of closely related
interacting partners.
Above: The top two panels represent two real networks describing the interactions
between plants (green nodes) and insects (orange nodes). The mutualist network (top
left) is a pollination community in Zackenberg, Greenland (Olesen et al. 2008), and
the antagonist network (top right) is a plant-herbivore community in the Espinhaço
mountain range, Southeast Brazil (Prado & Lewinsohn 2004). The bottom two panels
show interactions (blue squares) between plant and animal species; they are
demonstrative only and not based on real data.
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Methods S1
To compile our literature database, we searched for original research papers on the
effects of one or more global change drivers on one or more interspecific interaction
types. Note, the response variable of interest was the strength or frequency of the
interaction itself, rather than the diversity of species, the response of individual
species involved in the interaction, or the effects of an interaction on the response of a
species to a GEC driver. To focus our review, we used only empirical studies (not
theoretical papers, reviews or editorial material) of terrestrial systems. To augment the
studies with which the authors were familiar, we searched the ISI Web of Science
database for studies from 1997 to 2007 using the following search terms:
(rate or proportion or infect* or strength or dominance or advantage) and (pollinat*
or predat* or parasit* or pathogen or competit* or "food web" or hemiparasit* or
mycorrhiz* or endophyt* or herbivor* or (seed and dispers*)) and ("climate change"
or "elevated temperature" or drought or "nitrogen addition" or "elevated nitrogen" or
"invasive species" or "biotic invasion" or (habitat and (loss or degradation or
fragment*)) or ("land use" and (change or modif* or intensifi*)) or "CO2 enrichment"
or "elevated CO2" or "elevated carbon dioxide" or "carbon dioxide enrichment") not
(marine or aquatic or freshwater).
We further refined the search by excluding the subject areas:
(mathematical & computational biology or geography or materials science, ceramics
or mathematics, interdisciplinary applications or metallurgy & metallurgical
engineering or urban studies or food science & technology or limnology or
archaeology or marine & freshwater biology or cell biology or chemistry, organic or
engineering, environmental or computer science, artificial intelligence or computer
science, interdisciplinary applications or polymer science or computer science, theory
& methods or biochemistry & molecular biology or toxicology or engineering,
electrical & electronic or biophysics or geology or genetics & heredity or chemistry,
applied or health care sciences & services or materials science, multidisciplinary or
health policy & services or history & philosophy of science or geography, physical or
paleontology or instruments & instrumentation or materials science, composites or
oceanography or water resources or mathematics, applied or fisheries or chemistry,
analytical or nanoscience & nanotechnology or engineering, mechanical or obstetrics
& gynecology or pharmacology & pharmacy or physics, fluids & plasmas or
geosciences, multidisciplinary or nuclear science & technology or physics,
mathematical or anthropology or physics, multidisciplinary or developmental biology
or physics, nuclear or energy & fuels or psychology, biological or engineering,
chemical or spectroscopy or meteorology & atmospheric sciences or geochemistry &
geophysics or thermodynamics).
Of the approximately 1600 papers retrieved in this search, over 600 matched our
response variables of interest. These studies are presented as a summary spreadsheet
(Table S1) and were used to generate Fig.1 in the main text and Figs. S1-S6 in the
S.O.M..
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Figures S1-S6: Additional information supporting the trends in
changes to pairwise interactions depicted in main text Fig. 1
Predicted future changes to species interactions resulting from the effects of each
global environmental change (GEC) driver. Arrows with solid outlines indicate
nutrient and energy flow, while double-headed arrows with dotted outlines indicate
resource competition. + and – symbols within arrows indicate benefit or cost to each
participant (e.g. + + within an arrow is a mutualism). The proportion of colours within
each arrow indicates the proportion of interactions (from all interactions of a given
type and GEC driver present in our database Table S1) showing increases (green), no
effect (white), or decreases (dark grey) respectively in strength or frequency of the
interaction following each of five major GEC drivers. In Table S1, each row
represents the response of a single interaction type to a single global change driver. In
cases where a single published study examined effects of more than one driver or
responses of more than one interaction type, these results are presented in separate
rows. The number of specific pairwise interactions examined in the study (e.g. species
A interacts with species B) was recorded where possible; however, in some cases
responses were measured at the entire community level and individual pairwise
interaction responses could not be meaningfully separated. The number of these
pairwise interactions (or entire communities) showing a decrease, increase, or no
effect in the strength or frequency of the interaction, under that specific global change
driver was recorded, and this number of interactions was the basis for quantifying
changes to interaction frequencies in Fig. 1 and Figs. S1-S6. This essentially gave
extra weighting to studies that examined a greater variety of pairs of interacting
species. Of course we emphasise that this approach of vote counting can only give an
estimate of broad trends in the literature. Yellow arrows indicate a change in
dominance between competing species, which was recorded as a separate column in
Table S1. Width of arrows represents the number of studies considered in this review
(small: ≤ 10; medium: 11-40; large: > 40 cases) and in Fig. S6 the number of studies
on all drivers are combined (small: ≤ 40; medium: 41-100; large: > 100 cases). A
table of the studies on which these trends are based (Table S1) is provided in the
S.O.M, with details regarding specific treatments and response variables. Below each
supplementary figure we explain in more detail the potential effects of the driver on
each interaction, using a subset of references from our database (Table S1), and
relevant reviews or theoretical work (which were excluded from the database).
Although we acknowledge that this “vote-counting” approach can only give a very
approximate indication of trends in the literature, we believe that a quantitative metaanalysis of such a large number of different response variables and specific treatments
would give a false sense of confidence in the trends. We therefore emphasise that
these are broad generalizations based on a survey of original research studies from the
literature over the past decade and that many exceptions exist. The total trends for Fig.
1 in the main text and Fig. S6 are derived from the combination of the expected
changes in each of the individual drivers presented here. Figures S1-S6 are replicas of
the individual panels in main text Fig. 1. Roman numerals describe the interactions as
listed below each figure.
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Figure S1: CO2 enrichment
CO2 enrichment
Although increases in atmospheric CO2 concentrations can potentially drive climate
change, we separate these two drivers due to their differing effects on species and
their interactions. In particular, the effects of CO2 enrichment on plant growth may
contrast with the effects of climate change.
i)
Plant-pollinator: General benefit to pollinators, but variable. CO2
enrichment can induce changes to nectar quantity (increases; Lake & Hughes 1999;
Dag & Eisikowitch 2000; Davis 2003a; decreases; Rusterholz & Erhardt 1998; and no
change; Lake & Hughes 1999 have all been recorded) and composition (Erhardt et al.
2005), which can increase fecundity of, and attractiveness to, pollinator species
(Mevi-Schutz et al. 2003; Mevi-Schutz & Erhardt 2005). There is some variation in
these effects across plant species (Davis 2003a; Erhardt et al. 2005), and marginally
significant disruption to this mutualism can occur due to changes in flowering
phenology (Erhardt et al. 2005). The number of flowers produced has also been
shown to increase (Osborne et al. 1997) or remain unchanged (Lake & Hughes 1999)
by elevated CO2.
ii)
Plant-fungal: Frequent increase in colonisation, but highly variable.
Enrichment by CO2 may increase root colonisation by ectomycorrhizal fungi (EMF)
(Rey et al. 1997; Langley et al. 2003; Lukac et al. 2003) and arbuscular mycorrhizal
fungi (AMF) (Klironomos et al. 1998; Rillig & Allen 1998; Rouhier & Read 1998;
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Louche-Tessandier et al. 1999; Olesniewicz & Thomas 1999; Staddon et al. 1999;
Rillig et al. 2000; Hartwig et al. 2002; Hu et al. 2005) and increased extraradical
mycorrhizal hyphal production can occur (Gamper et al. 2004; Staddon et al. 2004),
but not always (Lewis et al. 1994; Kasurinen et al. 1999; Fransson et al. 2001; Jifon
et al. 2002). Increases may occur in subsoil but not topsoil (Rillig & Field 2003), and
only with coarse but not fine AMF endophytes (Rillig et al. 1999; Rillig & Field
2003). Elevated CO2 generally increases mycorrhizal fungal abundance (Treseder
2004) (possibly due to increased fungal sporulation; Klironomos et al. 1997b) and can
cause a shift in fungal strains (Gamper et al. 2005) and species (Parrent et al. 2006),
although other studies have also reported no significant effect on community
composition (Chung et al. 2006). Where they occur, shifts in fungal communities
have variable effects on plant growth, ranging from negative to positive, depending on
plant and fungal species (Rouhier & Read 1998; Gavito et al. 2000; Johnson et al.
2005c). Further, changes to fungal metabolism may have important consequences for
C cycling (Chung et al. 2006). The effect of elevated CO2 may be greater on EMF
than on AMF or plants (Alberton et al. 2005). Enrichment of CO2 may increase N
uptake (Hu et al. 2005), but no significant effect has been shown for P uptake
(Sanders et al. 1998; Staddon et al. 1998) or within-plant carbon allocation (Rouhier
& Read 1999). There is possible selection for fungal strains that help the host plant to
meet nutrient demands, and an indirect increase in N fixation by bacteria (Gamper et
al. 2005). Responses vary in strength or direction depending on the plant (Wolf et al.
2003; Gamper et al. 2005; Johnson et al. 2005c) and mycorrhizal (Rillig & Field
2003) species.
iii)
Seed dispersal: Unclear. To our knowledge no study to date has investigated
direct effects of CO2 on seed dispersal.
iv)
Plant-plant competition: Shift in advantage. Enrichment of CO2 may give a
competitive advantage to some plant species over others (Clark et al. 1997; Hebeisen
et al. 1997; Berntson et al. 1998; Diaz et al. 1998; Lovelock et al. 1998; Atkin et al.
1999; Navas et al. 1999; Greer et al. 2000; Huxman & Smith 2001; Reich et al. 2001;
Tilman & Lehman 2001; Fuhrer 2003; Poorter & Navas 2003; Zavaleta et al. 2003;
Stiling et al. 2004), through interspecific differences in the stimulation of growth
(Clark et al. 1997; Berntson et al. 1998; Diaz et al. 1998; Atkin et al. 1999; Greer et
al. 2000; Reich et al. 2001; Fuhrer 2003; Zavaleta et al. 2003) (e.g. C3 plants; Bazzaz
1990; Patterson et al. 1999 or fast-growing trees; Laurance et al. 2004) or
occasionally mediated by herbivore pressure (Diaz et al. 1998). ‘Mesic’ legumes
might be favoured over grasses and some Brassicaceae (Hebeisen et al. 1997;
Grunzweig & Korner 2001). No change was found in the relative competitive ability
of two C3 grasses (Hely & Roxburgh 2005), but invasive C4 grasses were found to
gain a competitive advantage over native C4 grasses due to increased germination and
growth rates (Baruch & Jackson 2005). Variation in responses can be due to nutrient
availability (Poorter & Navas 2003) or differences between species within functional
groups (Reich et al. 2001).
v)
Plant-hemiparasite: Frequent but variable benefit to parasite.
Hemiparasites may be affected directly through impacts on their physiology and
indirectly through impacts on host plants (Phoenix & Press 2005). The hemiparasite
may be favoured by higher growth response and increased carbon gains (Matthies &
Egli 1999; Grunzweig & Korner 2001; Hattenschwiler & Zumbrunn 2006),
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potentially increasing the demand for host mineral nutrients (Phoenix & Press 2005)
and competition with the host for N (Hwangbo et al. 2003). Alternatively, elevated
CO2 has been shown to have no effect (Watling & Press 1997, 1998; Matthies & Egli
1999) or even alleviate (Dale & Press 1998; Watling & Press 2000) the negative
effect of hemiparasites on their hosts and reduce the benefit for the parasite
(Grunzweig & Korner 2001). Variability can arise due to host species or nutrient
availability (Matthies & Egli 1999).
vi)
Plant-herbivore: Generally negative, but variable effects on herbivores.
Herbivore development times may increase (Johns & Hughes 2002; Asshoff &
Hättenschwiler 2005), decrease (Johns & Hughes 2002) or be unaffected (Awmack et
al. 2004; Chong et al. 2004) by CO2 enrichment. A recent meta-analysis (Stiling &
Cornelissen 2007) found a general decrease in herbivore abundance under elevated
CO2. Herbivore performance (Stiling et al. 1999; Johns & Hughes 2002; Percy et al.
2002; Veteli et al. 2002; Asshoff & Hättenschwiler 2005; Zvereva & Kozlov 2006),
and fecundity (Awmack et al. 2004; Asshoff & Hättenschwiler 2005) have been
shown to decrease, but may in some cases remain constant (Diaz et al. 1998; Bezemer
et al. 1999) or increase (Bezemer et al. 1999; Stacey & Fellowes 2002; Williams et al.
2003; Chen et al. 2005). Further, variation in within-plant physiological responses
may produce diverse responses of herbivores to CO2 enrichment (Pritchard et al.
2007). The varied responses of fecundity above give rise to variable effects on
herbivore population size (Docherty et al. 1997; Hughes & Bazzaz 2001; Stiling et al.
2002). Effects on oviposition choice and feeding preference were similarly found to
be variable (Docherty et al. 1997; Abrell et al. 2005; Hamilton et al. 2005).
Nevertheless, when it occurs, enhanced herbivore damage under elevated CO2 is
usually driven by elevated rates of overall consumption to compensate for reduced
food quality (Hughes & Bazzaz 1997; Stiling et al. 1999; Williams et al. 2000;
Bidart-Bouzat 2004; Handa et al. 2006), although numerous studies have found no
effect (Diaz et al. 1998; Peters et al. 2000; Williams et al. 2000; Johns et al. 2003;
Barbehenn et al. 2004; Cleland et al. 2006), reduced rates of consumption (Asshoff &
Hättenschwiler 2005; Cleland et al. 2006) or variation according to plant species
(Hattenschwiler & Schafellner 2004; Asshoff & Hättenschwiler 2005; Knepp et al.
2005). An early review (Bezemer & Jones 1998) suggested that leaf chewers such as
caterpillars can often compensate for reduced food quality by increasing their
consumption, but this was found not to be the case for grasshoppers (Asshoff &
Hättenschwiler 2005). Leafminers may (Stiling et al. 1999) or may not (Johns &
Hughes 2002) compensate for reduced plant quality, and phloem feeders even seem to
benefit from CO2 enrichment of the host plant (Bezemer & Jones 1998; Bezemer et
al. 1998; Chen et al. 2005). Protection against herbivores provided by mutualistic
endophytes has been found to increase under elevated CO2 (Marks & Lincoln 1996).
Effects of CO2 may be mediated by N fertilisation (Cleland et al. 2006), temperature
(Johns & Hughes 2002; Veteli et al. 2002; Zvereva & Kozlov 2006), host plant
species (Barbehenn et al. 2004) or ozone levels (Percy et al. 2002).
vii)
Plant-pathogen: Frequent increase in infection, but variable. Fungal
infection generally increases (Chakraborty et al. 2000; Mitchell et al. 2003), possibly
due to decreased water stress, increased leaf longevity or increased photosynthetic
rate (Mitchell et al. 2003). In addition to infection rates, the per capita effect of
pathogen infection on host photosystem II operating efficiency has been shown to
increase under elevated compared with ambient CO2 (Aldea et al. 2006). However,
10
there are exceptions (Chakraborty et al. 2000), with negligible effects on disease
expression reported in some instances (Meijer & Leuchtmann 2000). Moreover,
increased photosynthesis and water use efficiency, or the effects of CO2 concentration
on the transcription or post-translational turnover of pathogenesis-related proteins,
may lead to increased tolerance to fungal infection (Jwa & Walling 2001).
viii) Seed predation: Unclear. This has been seldom investigated, although one
study found predation of acorns by weevils to be unaffected by CO2 enrichment
(Stiling et al. 2004).
ix)
Parasite-host: Unclear. To our knowledge no study to date has investigated
direct effects of CO2 on parasite-host interactions.
x)
Animal-animal competition: Shift in advantage. Differences in benefit
among aphid species have been shown to alter competitive interactions (Stacey &
Fellowes 2002). See also variable effects on different herbivores in vi above, which
can cause a shift in competitive advantages.
xi)
Predator-prey or parasitoid-host: General increase in attack rates, but
variable.
Natural enemy densities have been shown to increase (Percy et al. 2002), and high
CO2 levels have been shown to enhance attack rates of herbivores by parastoids
(Stiling et al. 1999; Stiling et al. 2002). Prey consumption and mean relative growth
rates of predators may also increase (Chen et al. 2005). Where increased development
times of invertebrate herbivore larvae occur, this increases their period of maximum
vulnerability to predators and parasitoids (Johns & Hughes 2002; Asshoff &
Hättenschwiler 2005). In contrast, elevated CO2 may also reduce the volatile response
of plants to herbivory, making them less attractive to natural enemies of the
herbivores (Vuorinen et al. 2004). Other studies have shown no effect on
predation/parasitism rates (Bezemer et al. 1998; Stacey & Fellowes 2002; Awmack et
al. 2004), or found variation across species (Sanders et al. 2004) or plant genotypes
(Holton et al. 2003).
xii)
Decomposer food web: Some increases, but variable across all trophic
levels. Literature syntheses provide evidence that the soil microbial biomass (the basal
consumers of the decomposer food web) can show variable responses (Zak et al.
2000) (often positive (Phillips et al. 2002; Dijkstra et al. 2005; Hu et al. 2005;
Sonnemann & Wolters 2005) or neutral (Kandeler et al. 1998; Lussenhop et al. 1998;
Zak et al. 2000; Niklaus et al. 2001; Wiemken et al. 2001; Allen et al. 2005)
depending upon both plant species and nutrient availability (Klironomos et al. 1996).
Different effects on different species can also lead to shifts in the microbial
community composition (Phillips et al. 2002). Although only a handful of studies
have investigated effects of elevated CO2 on belowground consumers in higher
trophic levels, those that have show a variety of responses (Klironomos et al. 1996;
Lussenhop et al. 1998; Hungate et al. 2000; Wardle 2002; Yeates et al. 2003).
Nematodes can be positively (Li et al. 2007) or negatively (Neher et al. 2004)
affected, and their community structure can be altered (Hoeksema et al. 2000; Neher
et al. 2004; Li et al. 2007). Similarly, the microarthropod community has also been
shown to be positively (Jones et al. 1998; Sticht et al. 2006) or negatively (Niklaus et
al. 2003) affected, or to be unaffected (Klironomos et al. 1997a; Jones et al. 1998;
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Lussenhop et al. 1998). There are even instances in the literature in which higher level
consumers are promoted by CO2 enrichment but lower ones are not, which is
consistent with bottom-up regulation of organisms in higher trophic levels, which in
turn exert top-down effects regulating lower trophic levels (Lussenhop et al. 1998;
Yeates et al. 2003).
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Figure S2: N deposition
N deposition
i)
Plant-pollinator: Possible benefit to pollinators. If protein concentrations
increase in nectar as they do in leaves, attractiveness to pollinators could also increase
(Gardener & Gillman 2001; Mevi-Schutz & Erhardt 2005). Soil N can drive increases
in flower abundance, thereby attracting more pollinators, but this effect was found to
occur in the year following enhanced vegetative growth (Muñoz et al. 2005),
emphasising that N effects on plant-pollinator interactions may only be detectable at
long time scales. More research is required on this topic.
ii)
Plant-fungal: General decrease in colonisation by mycorrhizae.
Colonisation of grasses by non-mycorrhizal fungi increases following N deposition
(Siguenza et al. 2006), but effects on plant-mycorrhizal mutualisms are variable
(Hutchinson et al. 1998; Treseder 2004) (although often negative; Grogan & Chapin
2000; Treseder & Vitousek 2001; Hartwig et al. 2002; Staddon et al. 2004) across
taxa, with decreased AMF colonisation observed in native grasses but not legumes
(Gamper et al. 2004), and native shrubs but not exotic grasses (Siguenza et al. 2006).
Diversity and sporocarp abundance of aboveground (Lilleskov et al. 2001) and
belowground (Lilleskov et al. 2002) EMF, as well as EMF colonisation rates (Baum
& Makeschin 2000; Baum et al. 2002) have been shown to decline with increasing N.
There can also be a shift in EMF community structure (Avis & Charvat 2005), with a
decrease in fungi capable of using organic N sources, while certain “nitrophilic” taxa
13
were found to be unaffected or even benefit from increased N (Lilleskov et al. 2001).
Effects can also vary according to soil nutrient availability, with N addition causing
decreased allocation to AM structures at sites with ample P (i.e. low N:P ratio of soil),
and increased allocation to AM fungal structures when P is limiting (i.e. high N:P
ratio; Johnson et al. 2003).
Nitrogen deposition can negate the effects of increased CO2 on plantmycorrhizal mutualisms (West et al. 2005), and increasing soil nutrient availability
has been shown using modelling to cause a shift of plant-mycorrhizal interactions
from mutualistic to parasitic, as relative benefits to the plant decrease (Neuhauser &
Fargione 2004). Finally, increased N application can cause a decrease in the ratio of
fungal to total microbial biomass (de Vries et al. 2006), and soil acidification caused
by increased N can have negative effects on mycorrhizal communities (Bunemann et
al. 2006).
iii)
Seed dispersal: Possibly none. Although we are unaware of any research
directly testing the effects of N deposition on seed dispersal, low N and protein levels
in fruit (Jordano 2000), and the lack of variability in fruit morphology/chemistry
following N fertilization (Cipollini et al. 2004) suggest that large effects are unlikely.
iv)
Plant-plant competition: Shift in advantage. N deposition causes frequent
shifts in dominance (Hebeisen et al. 1997; Navas et al. 1999; Zavaleta et al. 2003)
due to advantages for certain N-demanding plant species (Bobbink et al. 1998;
Tilman & Lehman 2001; Brooks 2003; Rickey & Anderson 2004; Silliman &
Bertness 2004; Stevens et al. 2004; Badgery et al. 2005; Kuijper et al. 2005), with
competitive growth forms such as graminoids responding particularly well to soil
nutrient increases (Fuhrer 2003; Brooker 2006). Altered fungal infection may also
mediate competitive interactions between plant species, but other studies have found
no effect of N deposition on competition (Wilson & Tilman 1991; Strengbom et al.
2006). Effects may be mediated by management intensity (Hartley & Mitchell 2005).
v)
Plant-hemiparasite: Possibly none or slight decrease. Studies have shown
no effect of N on the plant-hemiparasite antagonism (Aflakpui et al. 2005) or a
decrease in hemiparasite seed germination with increasing N concentration up to a
point, then no further effect (Ayongwa et al. 2006).
vi)
Plant-herbivore: General increase in herbivory. N addition generally
makes plants more attractive to herbivores (Roy et al. 2004; Throop & Lerdau 2004;
Hines et al. 2005; Prudic et al. 2005; Cornelissen & Stiling 2006b) and enhances
herbivore population sizes (Armolaitis 1998; Fluckiger & Braun 1998; Haddad et al.
2000; Sudderth et al. 2005; Rowe et al. 2006), performance and consumption
(Kerslake et al. 1998; Nordin et al. 1998; Power et al. 1998; Hattenschwiler &
Schafellner 1999; Hartley et al. 2003; Throop & Lerdau 2004; Hartley & Mitchell
2005; Prudic et al. 2005; Stiling & Moon 2005; Throop 2005; Groenteman et al.
2006; Stevens & Jones 2006), although other studies have observed no effect on
consumption (Cleland et al. 2006), colony growth (Muller et al. 2005), and survival
(Cornelissen & Stiling 2006a), or even reduced consumption (Erelli et al. 1998) with
added N. Increased foliar concentrations of unpalatable or defence chemicals (Lou &
Baldwin 2004; Throop & Lerdau 2004) could make plants less attractive to
herbivores. Reduced carbon-based defence chemicals could also make plants
14
unattractive to herbivores that require those chemicals for their own defence (Prudic
et al. 2005). Increased production of alkaloids was found in endophyte-infested grass
following N addition, but this had no effect on aphids or their parasitoids (Krauss et
al. 2007). Variability has been observed between different interacting species (Roy et
al. 2004; Throop & Lerdau 2004), and effects may be most pronounced on low N
soils (Jiang & Schulthess 2005; Kuijper et al. 2005). Increased development rates of
herbivores (Groenteman et al. 2006) may offset the slowed development observed
under elevated CO2.
vii)
Plant-pathogen: General increase in infection. N deposition can cause
increased severity of pathogen infection (Nordin et al. 1998; Limpens et al. 2003;
Mitchell et al. 2003; El-Hajj et al. 2004; Strengbom et al. 2006), and stimulate
expansion of leaf-surface algae, reducing the volume of host photosynthetic tissue
(Limpens et al. 2003). Increased leaf amino acid concentrations can also promote
fungal infection (Strengbom et al. 2002), but high doses of N can increase production
of phenolics that may defend against fungal pathogens (Witzell & Shevtsova 2004).
Effects of N can be mediated by climatic factors (Strengbom et al. 2006).
viii) Seed predation: Variable, but possible decrease. Studies have shown
reduced abundance and effect of seed-head weevils in fertilized treatments (Lejeune
et al. 2005), no correlation between seed preferences of rodents and nitrogen
concentration (Kollmann et al. 1998), and increased availability of plant species of
value to granivorous birds (e.g., Stellaria - chickweeds) (Wilson et al. 1999) under
elevated N.
ix)
Parasite-host: Possible reduction in infection. Little is known, although
increased dietary protein from plants under elevated N has been shown to reduce
gastrointestinal nematode infection of small ruminants (Knox et al. 2006).
x)
Animal-animal competition: Shift in advantage. Different responses among
invertebrate herbivores to N addition have been shown (Roy et al. 2004; Throop &
Lerdau 2004), leading to shifts in dominance (Hines et al. 2005). Hares were also
found to show a preference for high N plants, although they avoided these plots when
geese (which also preferred high N plots) were present (Stahl et al. 2006).
xi)
Predator-prey or parasitoid-host: General increase in attack, but
variable. N fertilisation has been shown to lead to increased attack rates by
parasitoids (Moon et al. 2000; Moon & Stiling 2000). In contrast, shorter larval
development times may reduce the window of maximum vulnerability of insect
herbivores to predators and parasitoids (Mevi-Schutz et al. 2003; Cornelissen &
Stiling 2006a), and N fertilization was found to have no significant effect on leaf
miner mortality caused by natural enemies (Cornelissen & Stiling 2006a). Reduced
allocation of carbon to C-based structures such as trichomes may negatively affect
herbivore defences (Throop & Lerdau 2004), as may reduced larval defence
chemicals (Prudic et al. 2005). Finally, N deposition has been shown to lead to
increased predator to herbivore ratios (Hines et al. 2005), increased parasitoid and
hyperparasitoid abundance (Krauss et al. 2007), and increased egg load of parasitoid
offspring (Jiang & Schulthess 2005).
15
xii)
Decomposer food web: Variable across all trophic levels. Available
evidence points to the soil microbial biomass (primary consumer of the soil food web)
and the decomposition processes that it drives, showing positive (Johnson et al. 1998;
Lussenhop et al. 1998; Ruess et al. 1999; Bradley et al. 2006; Manning et al. 2006;
Power et al. 2006; Rinnan et al. 2007), neutral (Johnson et al. 1998; Wiemken et al.
2001; Dijkstra et al. 2005; Johnson et al. 2005a) or negative (Johnson et al. 1998;
Fisk & Fahey 2001) responses to N addition depending on context (Soderstrom et al.
1983; Kaye & Hart 1997; Scheu & Schaefer 1998; Bardgett et al. 1999b; Ettema et al.
1999). These variable effects are also propagated through higher trophic levels in the
soil food web, affecting flows through the fungal vs. bacterial energy channels
(Bardgett et al. 1999b; Ettema et al. 1999). Further, there is evidence for N addition
sometimes promoting higher trophic levels (e.g. nematodes; Ruess et al. 1999; Li et
al. 2007 and microarthropods; Klironomos et al. 1997a) in the soil food web but not
lower ones, presumably because of regulation of lower trophic levels by increased
predation (Ettema et al. 1999). However, Collembola may increase (Manning et al.
2006) or decrease (Sticht et al. 2006) in abundance.
16
Figure S3: Climate change
Climate change
To assess impacts of climate change we include studies that test effects of increase in
average temperature, altered rainfall regimes and increased frequency of extreme
weather events such as El Niño southern oscillations (ENSO).
i)
Plant-pollinator: General reduction in pollination. Drought-linked
reduction in flower availability can lead to extinction of specialist pollinators
(Harrison 2000, 2001), and interannual shifts in pollinator communities have been
correlated with climatic changes (Wall et al. 2003). Climate-induced changes to
flowering phenology (Price & Waser 1998; Fitter & Fitter 2002; Schauber et al. 2002;
Lambrecht et al. 2007) or abundance (Saavedra et al. 2003) can reduce temporal
overlap between plants and pollinators (Memmott et al. 2007), with timing and
intensity of masting events being particularly vulnerable to climatic changes
(Schauber et al. 2002). The quantity and composition of nectar produced may also be
affected by temperature and water availability (Pacini et al. 2003; Petanidou 2003),
affecting attractiveness to pollinators. Variation in response due to habitat context
(Kudo & Hirao 2006) has been found.
ii)
Plant-fungal: Highly variable. Increased root colonisation (Gavito et al.
2003; Staddon et al. 2004), extraradical mycorrhizal hyphal (Staddon et al. 2004) and
mycelial production (Clemmensen et al. 2006) have been shown with soil warming,
as have increased frequency of endophyte infection (Ju et al. 2006) and defence
17
alkaloid production (Salminen et al. 2005). However, responses of plant-mycorrhizal
mutualisms to climate change have been highly variable across plant species
(Heinemeyer & Fitter 2004), and often difficult to separate from climate effects on
host-plant physiology (Staddon et al. 2002), as colonisation may reflect increases in
plant biomass (Heinemeyer & Fitter 2004). Drought has been shown to cause
decreased colonisation by a fine endophyte (Staddon et al. 2004). Dark respiration in
lichens (algal-fungal associations) has been shown to acclimate to seasonal
temperature fluctuations (Lange & Green 2005).
iii)
Seed dispersal: General reduction in dispersal. Inadequate pollination may
have cascading effects on frugivorous vertebrates (Harrison 2000) that disperse seeds.
Shifts in seasonal fruit availability may disrupt the match between production peaks
of fruits and arrival of migratory or seasonal birds (Jordano 2000). Climate change
may also alter the primary agent of seed dispersal, due, for example, to different
effects on ants vs. rodents (Ness & Bressmer 2005).
iv)
Plant-plant competition: Shift in advantage. Climate change is well known
to cause major changes in vegetation composition and species ranges (Laurance et al.
2004; Brooker 2006; Parmesan 2006), and may give a competitive advantage to
certain species (Tilman & Lehman 2001; Zavaleta et al. 2003; Klanderud 2005; Wang
et al. 2006), such as C4 plants (Fuhrer 2003) or graminoids (Brooker 2006; Walker et
al. 2006). Changes to plant phenology brought about by global warming (Root et al.
2003) may affect the ability of different species to acquire resources early in the
season (Dunnett & Grime 1999), or shift herbivore preference (Russell & Louda
2005), thereby providing associational resistance. Time of snowmelt and associated
nutrient releases can alter community dominance (Heegaard & Vandvik 2004) and
productivity (Wasley et al. 2006). No change in competitiveness between two C3
grasses (Hely & Roxburgh 2005) implies that differences between major growth
forms may be more important than differences within groups. Climate change may
also increase the frequency of droughts in certain areas, which increases fire risk and
may interact synergistically with habitat clearance and burning to affect forest tree
species (Laurance & Williamson 2001).
v)
Plant-hemiparasite: Variable. Increased rates of photosynthesis may
increase the demand of hemiparasites for host mineral nutrients (Phoenix & Press
2005). The parasitic annual life style of a hemiparasite, without a persistent seed bank,
was found to make it vulnerable to spring drought, which induced population
collapses (Ameloot et al. 2006). Mineral nutrients may alleviate the impacts of
climate change on plant-hemiparasite associations (Phoenix & Press 2005).
vi)
Plant-herbivore: Frequent increase in herbivory, but highly variable.
Temperature may be the dominant abiotic factor directly affecting herbivorous insects
(Bale et al. 2002), and climate change may drive increased consumption (Johns et al.
2003), and shifts in species composition (Fuhrer 2003; Roy et al. 2004; Andrew &
Hughes 2005), host plant preference (Russell & Louda 2005) or range (Battisti et al.
2005). The performance (Johns & Hughes 2002; Veteli et al. 2002; Zvereva &
Kozlov 2006), survival (Kiritani 2007) and abundance (Bezemer et al. 1998; Logan et
al. 2003) of herbivores tends to improve under conditions of simulated climate
change, and development rates are also accelerated (Johns & Hughes 2002; Williams
et al. 2003; Chong et al. 2004), although variation in the response of different
18
components of plant physiology may produce diverse responses of herbivores to
drought (Pritchard et al. 2007). Mistiming due to phenological differences between
herbivores and their food plant may interfere with this antagonism (Visser &
Holleman 2001; Visser & Both 2005; Musolin 2007), but other studies have shown no
shifts in the phenological match between plant and herbivore (Sparks & Yates 1997).
There are also negative effects on herbivory, due to increases in endophytic toxins
affecting mammals (Ju et al. 2006) and insects (Salminen et al. 2005), and variation
in the effects of climate has been observed among different herbivore species (Roy et
al. 2004), with some species showing reduced growth and consumption (Williams et
al. 2000).
vii)
Plant-pathogen: General increase in infection. Many pathogens of plants
and animals are limited by climatic requirements for overwintering (Pfender &
Vollmer 1999; Garrett et al. 2006), and diseases may increase in incidence
(Strengbom et al. 2006) and expand in geographic range (Kamata et al. 2002; Fuhrer
2003; Roy et al. 2004; Parmesan 2006) due to climate warming, provided that
maximum threshold temperatures are not exceeded (Stacey 2003). Disease vectors
may also carry more infections with increasing temperatures (Fabre et al. 2005);
however, no effect of early snowmelt on pathogen occurrence has been observed in
some species (Roy et al. 2004).
viii) Seed predation: Variable, depending on direction of change in plant
phenology. Much research shows that the synchrony of insect activity with plant
resources can affect the impact of floral herbivores on their host plant populations
(Russell & Louda 2004). Changes in the timing of seed development may therefore be
crucial, as late opening of cones can lead to an increase in pre-dispersal seed
predation (Worthy et al. 2006). Early flowering/fruiting has been shown to either
significantly reduce seed predation by grasshoppers (Lacey et al. 2003), or increase
the severity of insect attack in other cases (Mahoro 2003). Climate change can also
alter the relative collection of seeds by ant dispersers or rodent predators (Ness &
Bressmer 2005). Increased average temperature and increasing variability in
temperature can even have contrasting effects, with the former causing an increase,
and the latter causing a decrease in pre-dispersal seed predation in grasslands
(McKone et al. 1998).
ix)
Parasite-host: Increase in infection. ENSO events have been shown to
promote the growth of disease vector populations (Stapp et al. 2004), and facilitation
of pathogen outbreaks by temperature shifts has been implicated in widespread
amphibian extinctions (Pounds et al. 2006). Climate change may facilitate arbovirus
spread (Toussaint et al. 2006) and transmission of nematode parasites between
mammals (Kutz et al. 2005) and birds (Cattadori et al. 2005). It has also been shown
to cause a population collapse of trematode-infected amphipods (Mouritsen et al.
2005). Increased temperatures may allow disease vectors (e.g. ticks; Lindgren &
Gustafson 2001) and parasites (Poulin & Mouritsen 2006) to increase in abundance.
Diseases may also expand their geographic range (Ebi et al. 2005; Ogden et al. 2006),
but the effects of climate may vary across regions (latitude, lowland vs. highland)
(Ebi et al. 2005).
x)
Animal-animal competition: Shifts in competitive ability. Climate change
may drive shifts in invertebrate species composition (Fuhrer 2003; Roy et al. 2004;
19
Andrew & Hughes 2005; Helms & Vinson 2005; Hodkinson 2005) and food web
structure (Polis et al. 1997), due to variation in the effects of climate on different
species (Roy et al. 2004). Invertebrates may sometimes benefit over vertebrates. For
example, increased temperature reduces carcass scavenging by vertebrates, but
increases activity by insects (DeVault et al. 2004). Further, temperature affects
dehiscence of seeds, which affects competition between rodent seed predators and ant
dispersers (Ness & Bressmer 2005), and changes in global temperature and rainfall
may create gradients of ectotherm size and affect competition between small and large
species (Kaspari 2005). The importance of competitive interactions among small
mammals was also shown to vary greatly with changing local demography, which
was driven largely by climatic patterns (Kelt et al. 2004).
xi)
Predator-prey or parasitoid-host: Possible increase, but highly variable.
Higher trophic levels are likely to be more susceptible to climate change and will be
disproportionately lost from communities (Petchey et al. 1999; Voigt et al. 2003).
Climate change can affect the timing (Visser et al. 2003; Both et al. 2006),
demography (Kelt et al. 2004) and abundance (Durant et al. 2003) of prey species,
thereby disrupting predator-prey dynamics (Durant et al. 2005; Visser & Both 2005).
Shorter development times of insect herbivore larvae can reduce vulnerability to
predators and parasitoids (Johns & Hughes 2002), and parasitism rates have been
shown to decline with increasing climatic variability (Stireman et al. 2005). Parasitoid
longevity (Chong & Oetting 2006), development inside the host (Fellowes et al. 1999;
Hegazi & Khafagi 2005) and rates of predation and parasitism (Ris et al. 2004) have
all been shown to decline with increasing temperature. In contrast, other studies have
shown increased development rates (Chong et al. 2005), consumption rates (Skirvin et
al. 1997; Bezemer et al. 1998; Perdikis et al. 1999; Virtanen & Neuvonen 1999; Polis
et al. 2000; Van Nouhuys & Lei 2004; Martin 2007), and abundance (Roy et al. 2004)
of predators and parasitoids. Moisture associated with El Niño events has been shown
to facilitate top-down control of herbivores by increasing the persistence of their
nematode predators (Preisser & Strong 2004). Foraging efficiency and consumption
rates of insect predators has been found to increase more than that of their prey
following temperature increases (Stacey 2003), and parasitoids can also be favoured
by warm early spring temperatures through effects on host-parasitoid synchrony (Van
Nouhuys & Lei 2004). Additionally, climate change may indirectly facilitate predatorprey interactions by altering vegetation structure and thereby affecting prey
susceptibility to predators (Martin 2007).
xii)
Decomposer food web: Variable across all trophic levels. Elevated
temperature has effects on the soil microbial biomass that are positive (Ruess et al.
1999), neutral (Bardgett et al. 1999a; Ruess et al. 1999; Zhang et al. 2005), or
negative (Arnold et al. 1999; Waldrop & Firestone 2006; Rinnan et al. 2007)
depending on context (Kandeler et al. 1998). Similar results have been found with
regard to the community structure of soil invertebrates occupying higher trophic
levels (Cole et al. 2002; Convey et al. 2002; Dollery et al. 2006), indicative of
bottom-up regulation of higher level consumers (Harte et al. 1996; Briones et al.
1997; Ruess et al. 1999; Sohlenius & Boström 1999).
20
Figure S4: Biotic invasions
Biotic invasions
Although there has been a strong theoretical focus on the importance of species
interactions as determinants of invasion success (Mitchell et al. 2006), there has not
been a comparable research focus on the resulting impacts of invasive species on
species interactions within invaded communities (White et al. 2006). Unlike other
GEC drivers which can have direct effects on pairwise species interactions, biotic
exchange involves embedding a novel species into an existing set of species
interactions, so that strictly speaking any impact of an invasive species on the pairwise interaction between two other species would classically be referred to as an
indirect effect within food web ecology. Consequently, we do not focus on the many
examples of direct effects of an invasive species on the phenotype or abundance of a
native species, but rather on the indirect effects of invasive species on ecological
interactions between other species (native or non-native). In Figure S4, invasive
species are depicted within circles, and the new interactions between invasive and
native biota are depicted as an interaction (block arrow) with increasing strength.
These new links are added for consistency, but the Roman numeral footnotes refer to
indirect effects of biotic exchange on already existing interactions.
i)
Plant-pollinator: General disruption of pollination. Growing evidence now
supports two primary mechanisms of indirect disruption to native plant-pollinator
mutualisms: through exploitative or interference competition for available flower
resources by invasive pollinators (depicted symbolically in Figure S4 as a honeybee)
21
(Gross & Mackay 1998; Kato et al. 1999; Gross 2001; Hansen et al. 2002; Celebrezze
& Paton 2004; Ings et al. 2006), and through exploitative competition for available
native pollinators by invasive plants (Chittka & Schurkens 2001; Brown et al. 2002;
Ghazoul 2004; Moragues & Traveset 2005). The latter may lead to reduced
conspecific pollen on stigmas of native plants in plant-invaded plots (Larson et al.
2006), but alternatively, native pollinators may be somewhat averse to feeding on
exotic plants, possibly due to their rarity (Memmott & Waser 2002). Despite these
effects on native pollinators, exotic plants can be sufficiently pollinated by exotic bees
when suitable native pollinators are absent (Stout et al. 2002). In addition, a few
studies show that indirect antagonistic interactions at higher trophic levels may also
impact on plant-pollinator mutualisms, with comparatively strong evidence for
invasive predators (depicted symbolically in Figure S4 as a mustelid) altering the
abundance of native bird and lizard pollinators (Traveset & Saez 1997; Kelly et al.
2006), or invasive ants reducing pollination of native plants (Blancafort & Gomez
2005). Only limited evidence exists so far for indirect negative effects of invasive
herbivores on plant-pollinator interactions (Vazquez & Simberloff 2004; Traveset &
Richardson 2006), possibly through effects on plant and pollinator population growth
(Spurr & Anderson 2004).
ii)
Plant-fungal: General but variable reduction in colonisation. Invasive
plants (Batten et al. 2006; Mummey & Rillig 2006) and earthworms (McLean et al.
2006) may affect AMF community composition, diversity and abundance, and reduce
extraradical hyphal lengths, but this may (Klein et al. 2006) or may not (Pritekel et al.
2006) translate into differences in AMF infection intensity between plant-invaded vs.
uninvaded communities. Mycorrhizal inocula from invaded grasslands may also
improve plant growth relative to native grassland inocula (Gillespie & Allen 2006).
However, interactions between invasive plants and AMF communities may facilitate
further invasion by using antifungal phytochemicals to disrupt the mutualism and
native plant growth (Stinson et al. 2006) or by parasitizing mycelial networks
between multiple plants to allow the invasive to establish initially and then reduce
AMF availability (Reinhart & Callaway 2006). Fungi themselves may also potentially
become invasive (Schwartz et al. 2006). For example, exotic EMF have been shown
to facilitate the spread of exotic trees (Diez 2005), and a toxic fungal endophyte
introduced with the grass Lolium arundinaceum altered rates of sympatric tree
herbivory and slowed succession from grassland to forest (Rudgers et al. 2007).
iii)
Seed dispersal: General disruption of dispersal. Although there has been
substantially less research emphasis placed on seed dispersal than on pollination,
species invasions are thought to disrupt both mutualisms through substantively the
same mechanisms (Traveset & Richardson 2006). Invasive seed dispersers can reduce
the number of fruit visits or seeds removed by native dispersers (Ferguson & Drake
1999; Christian 2001; Carney et al. 2003). For example, Traveset and Riera (Traveset
& Riera 2005) documented the disruption of a strong interaction between an endemic
perennial shrub and an endemic frugivorous lizard, precipitated by the displacement
of the lizard by introduced mammals since Roman times. Invasive species may also
promote seed deposition in suboptimal germination sites (Riera et al. 2002), and alter
the frequency of fruit predation rather than seed dispersal (Kelly et al. 2006). For
example, invasive ants may displace important seed dispersers (Witt & Giliomee
2004), and this can lead to reduced seed dispersal distances (Ness et al. 2004). Some
examples of invasion-induced disruption of seed dispersal mutualisms have led to
22
cascading effects on plant community assembly over moderately large spatial scales
(Christian 2001).
iv)
Plant-plant competition: Shift in advantage. Herbivory by invasive insects
and mammals can indirectly facilitate dominance of invasive plants over natives
(Schierenbeck et al. 1994; Callaway et al. 1999). Compensatory growth, changes to
the mycorrhizal community, or harmful root exudates produced as invasive plants
respond to herbivory may have deleterious effects on native plants (Pearson &
Callaway 2003). Similarly, Edwards and others (Edwards et al. 2000) found that
invasive rabbits (Oryctolagus cuniculus) promoted a shift in competitive balance
between the invasive creeping thistle (Cirsium arvense) and several native grass
species, allowing creeping thistle to dominate. Invasive deer browsing has also been
found to promote native bryophyte growth, altering the relative competitive advantage
of invasive Rhododendron over native shrubs (Cross 1981). Competition in intertidal
kelp communities can be altered by an invasive epiphytic alga, which causes a
competitive reversal favouring an invasive alga over native kelp (Levin et al. 2002).
Finally, invasive pathogens have been shown to preferentially attack native grasses,
causing a competitive reversal in favour of invasive grasses (Malmstrom et al. 2006).
v)
Plant-hemiparasite: Unclear. To our knowledge no study to date has
investigated effects of invasive species on hemiparasite-host interactions.
vi)
Plant-herbivore: Frequent, but variable increase in herbivory. Invasive
herbivores can have strong negative effects on native (Franks et al. 2006) and
invasive (Halpern & Underwood 2006; Liu & Stiling 2006) plants. In many situations
invasive herbivores can cause changes in competitive balance between native and
invasive plants (see iv above), and in turn some invasive plants facilitate an increased
negative impact of invasive herbivores on native plants. For example, Rand and
Louda (2004) found that exotic thistle (Carduus nutans) invasion increased the
susceptibility of native Cirsium undulatum plants to an invasive biocontrol weevil
(Rhinocyllus conicus). Similar examples of invasive herbivore-mediated apparent
competition have been shown to increase attack rates on native plants in the presence
of invasives, in New Zealand (Sessions & Kelly 2002), California (Lau & Strauss
2005), and Hawaii (Lenz & Taylor 2001). Invasive ants can also promote attack of
native plants by exotic herbivores (O'Dowd et al. 2003). In contrast, invasive
herbivores can outcompete natives for shared resources, causing a reduction in native
herbivore growth/survival (Byers & Noonburg 2003). Salt marsh invasions have been
shown to cause a shift from plants to detritus being used as the basal resource in
arthropod webs (Gratton & Denno 2006). Native plant-herbivore interactions are
further altered by species invasions at higher trophic levels, through the cascading
effects on native and invasive herbivores (see below).
vii)
Plant-pathogen: Possible increase in infection. Exotic plants can indirectly
influence virus incidence in native plants by increasing populations of the aphid
vector (Malmstrom et al. 2005). Exotic plants may potentially also act as source
populations for pathogens themselves (Dwyer et al. 2007).
viii) Seed predation: General, though not universal, increase in predation.
Invasive seed predators (depicted symbolically in Figure S4 as an ant) can have
stronger effects than native seed predators on native plants, potentially driving
23
extinction of the plant and indirectly affecting its endemic seed predators (Rose et al.
2005). Reduced seed dispersal in areas invaded by ants was found to lead to a higher
proportion of seeds being predated by rodents (Witt & Giliomee 2004). Plant
invasions can also affect seed predation. For example, exotic weeds were found to
provide an important food source for sustaining endangered native granivore
populations (Schiffman 1994), but attack on native thistles increased strongly with
increasing density of exotic thistles (Rand & Louda 2004). A further study found that
granivores removed an order of magnitude less native seeds than exotic seeds
(Folgarait & Sala 2002).
ix)
Parasite-host: General increase in transmission. Invasive hosts can carry
new parasites with the potential to remain in a system even after their founding host is
extirpated (Smith & Carpenter 2006), although a similar number of blood parasite
lineages was found in native populations compared to introduced populations (Ishtiaq
et al. 2006). Invasive species may act as vectors (Lilley et al. 1997; Kiesecker et al.
2001; Tompkins & Gleeson 2006) or source populations (Tompkins et al. 2002;
Tompkins et al. 2003; Hampton et al. 2004) for diseases, increasing spread among
native hosts. In contrast, an invasive species that is a poor quality host has been
shown to have a dilution effect, reducing parasite infection of native hosts (Telfer et
al. 2005).
x)
Animal-animal competition: Shift in advantage. While a large body of
literature has demonstrated direct competitive effects of invasives on natives (outside
the scope of this review), there are comparatively few examples of invasive species
altering competitive interactions between other species. However, displacement of
native ants by invasive species can potentially alter food web structure (Rowles &
O'Dowd 2007), and mutualisms between invasive ants and aphids have been shown to
reduce the survival of herbivore competitors. In Hawaii, the invasive gecko
Hemidactylus frenatus displaced the native Lepidodactylus lugubris indirectly through
more effective exploitation of available insect prey, rather than by direct interference
competition (Petren & Case 1996). In at least two cases, an invasive species has been
shown to affect native competitors by apparent competition mediated through
increased density of a shared natural enemy (a native parasitoid of native leafhoppers;
Settle & Wilson 1990, or a shared predatory crab feeding on native clams; Grosholz
2005). However, invasive mammals (McDonald et al. 2007) and ants (King &
Tschinkel 2006) may not always outcompete natives, but rather increase in abundance
when the native declines due to an external driver. Further, numerical dominance by
invasive ants may be greatest immediately following invasion, then decline over time
(Morrison 2002; Strayer et al. 2006). Although exotic species may be competitively
superior (Turnock et al. 2003; Yasuda et al. 2004), the same has been shown for
natives, which may even act as intraguild predators (Hickerson et al. 2005).
Compared to trophic interactions, competition from introduced species is not likely to
be a common cause of extinctions of long-term resident species (Davis 2003b). For
example, invasive starlings do not have severe impacts on populations of native birds
(Koenig 2003). Competitive exclusion of native earthworms by exotic earthworms is
not easily demonstrated and, in fact, co-existence of native and exotic species appears
to be common, even if transient (Hendrix et al. 2006).
xi)
Predator-prey or parasitoid-host: General but variable reduction in
predation. Introduced generalist predators can have particularly large effects on
24
native species (Johnson et al. 2005b; Snyder & Evans 2006), even altering the
primary energy channels used (Geiger et al. 2005). In Delaware (USA) old fields,
invasive Chinese mantis (Tenodera sinensis) displaces native spider predators, but
also preys directly on native herbivores, with the net effect that herbivore abundance
decreases, leading to enhanced plant growth (Moran et al. 1996). Displacement or
predation of native predators by invasives is also observed in coccinelid beetles
(Alyokhin & Sewell 2004; Evans 2004), intertidal crustaceans (Grosholz et al. 2000),
and lizards (Suarez & Case 2002) Changes in basal resource availability due to plant
invasion of wetlands can alter the structure of arthropod food webs (Gratton & Denno
2006) and an invasive plant and its herbivorous biocontrol agent have been shown to
provide an effective resource subsidy that increases the relative abundance of native
parasitoids in heavily-invaded habitats (Willis & Memmott 2005). Such subsidies of
invasive prey can increase consumer abundance and lead to spillover of consumers
onto native prey (Settle & Wilson 1990; Pearson et al. 2000; Norbury 2001; Benson
et al. 2003; Ortega et al. 2004; Noonburg & Byers 2005; Rand et al. 2006). Feral pigs
have been shown to provide a similar resource subsidy to golden eagles, which then
drive population declines of an endemic fox (Roemer et al. 2001; Courchamp et al.
2003), but reduce predation rates of foxes on native skunks (Roemer et al. 2002).
Conversely, an invasive ladybird has been shown to act as a population sink for native
parasitoid eggs, resulting in a population increase of native ladybirds (Hoogendoorn
& Heimpel 2002). Additionally, an invasive plant can affect conspicuousness of
(lizard) prey to predators, thereby altering predation rates (Valentine et al. 2007).
Introduced prey can also drive evolution of predator behaviour and physiology
(Phillips & Shine 2006). In an extreme case, a toxic invasive pest kills its predators
(Suttle & Hoddle 2006), possibly releasing other prey from predation pressure. In
other cases, interference between introduced predator species (Snyder & Evans 2006)
may reduce predation on their shared prey (Griffen & Byers 2006). Predators can
favour introduced over endemic prey species (Griswold & Lounibos 2005), but this
does not necessarily lead to reduced consumption of native prey (Maerz et al. 2005).
Positive and negative effects may even occur simultaneously. For example, in alfalfa
fields in Utah (USA), an invasive predator (seven-spot ladybird, Coccinella
septempunctata) suppresses weevil abundance through predation, and simultaneously
enhances weevil survival by eating the aphids that provide resource subsidies
(honeydew) to weevil parasitoids (Evans & England 1996). Introduced parasitoids can
form new interactions with exotic and native hosts (Elkinton et al. 2006), but at least
for specialist parasitoids the overall impact on herbivore populations is less than
generalist predators, and usually not enough to cause population extinction (Johnson
et al. 2005b; Keeler et al. 2006).
xii)
Decomposer food web: Variable (depends on type of invader): Invasive
plant species often (though not always; Belnap et al. 2005) promote decomposer
microbes and the processes that they drive (Burtelow et al. 1998; Saggar et al. 1999;
Ehrenfeld 2003; Van der Putten et al. 2007), and these effects can propagate through
to higher trophic levels (Yeates & Williams 2001; Standish 2004). Shifts in
dominance within the microbial (Kourtev et al. 2003; Marschner et al. 2005; Batten et
al. 2006; Li et al. 2006) and microfaunal (Yeates & Williams 2001) community are
also commonly observed in the presence of invasive plants. In addition, invasive
aboveground consumers can greatly affect decomposer biota, but the nature of effect
is dependent upon the context, including the type of invader involved (Wardle et al.
2001; Fukami et al. 2006). Invasive earthworms may promote microbial activity
25
(Bohlen et al. 2004) but can have a range of effects on both soil microbes and soil
fauna (McLean & Leckie 2000), causing declines and shifts in dominance of
macrofaunal communities (Chauvel et al. 1999), microarthropods (McLean &
Parkinson 1998a, 2000) and soil fungi (McLean & Parkinson 1998b, 2000). Invasive
predatory flatworms can significantly reduce earthworm densities and therefore the
effects that earthworms have on other biota (Boag 2000). Soil invertebrates have also
been shown to be negatively affected by invasive ants (Gotelli & Arnett 2000), beetles
(Niemala et al. 1997), and grass (Gremmen et al. 1998).
26
Figure S5: Land use change
Land use change
We use the broad term “land use change” to encompass all anthropogenic changes to
the abundance and structure of natural habitats, including habitat loss, fragmentation,
increased agricultural management intensity, and altered abiotic (e.g., hydrological)
and biotic (e.g., grazing) disturbance regimes.
i)
Plant-pollinator: General reduction in pollination. The large body of work
examining the effects of land use changes on plant-pollinator mutualisms has revealed
predominantly negative responses of pollinators and pollination success to habitat
fragmentation (Jennersten 1988; Lamont et al. 1993; Oostermeijer & van Swaay
1998; Bigger 1999; Gigord et al. 1999; Jules & Rathcke 1999; Morgan 1999; SteffanDewenter & Tscharntke 1999; Luijten et al. 2000; Nielsen & Ims 2000; Parra-Tabla et
al. 2000; Somanathan & Borges 2000; Warburton et al. 2000; Ghazoul & McLeish
2001; Groom 2001; Moody-Weis & Heywood 2001; Mustajarvi et al. 2001; Rocha &
Aguilar 2001; Steffan-Dewenter et al. 2001; Wolf & Harrison 2001; Bosch et al.
2002; Bruna & Kress 2002; Gathmann & Tscharntke 2002; Jacquemyn et al. 2002;
Kremen et al. 2002; Leimu & Syrjanen 2002; Lennartsson 2002; Paschke et al. 2002;
Tomimatsu & Ohara 2002; van Rossum et al. 2002; Lienert & Fischer 2003; Quesada
et al. 2003; Severns 2003; Smith-Ramirez & Armesto 2003; Brys et al. 2004; Duncan
et al. 2004; Johnson et al. 2004a; Quesada et al. 2004; Rossetto et al. 2004; Blanche
& Cunningham 2005; Ghazoul 2005; Honnay et al. 2005; Kolb 2005; Ward &
27
Johnson 2005; Aguilar et al. 2006; Valdivia et al. 2006; Klein et al. 2007; Ockinger &
Smith 2007) and management intensity (Gabriel & Tscharntke 2007), but some
variability due to self compatible species being less affected than self incompatible
species (Aguilar et al. 2006). There have also been a few cases of increased pollinator
abundance or pollination in fragmented or modified habitats (Karrenberg & Jensen
2000; Kelly et al. 2000; Schmidt & Jensen 2000; Mavraganis & Eckert 2001; Johnson
et al. 2004b; Yates & Ladd 2005; Tylianakis et al. 2006; Diekotter et al. 2007).
Reduced pollination in fragmented habitats may lead to declines of animal-pollinated
tree species (Laurance et al. 2006), but the presence of corridors connecting
fragmented habitats may lead to increased pollination (Tewksbury et al. 2002).
Disruption to this mutualism is usually attributed to reduced pollinator diversity and
abundance (Rathcke & Jules 1993; Klein et al. 2003a; Tylianakis et al. 2005; Chacoff
& Aizen 2006), altered pollinator behaviour (Thompson 2001; Montgomery et al.
2003; Cheptou & Avendano 2006), or shifts in pollen quality transferred by different
species (Chacoff et al. 2008). Fragmentation can eventually lead to genetic drift
(Hooftman et al. 2004), and allee and inbreeding effects in plant (Galeuchet et al.
2005; Cheptou & Avendano 2006; Lazaro et al. 2006; Wagenius et al. 2007) and
pollinator (Darvill et al. 2006; Ellis et al. 2006) populations. However, variation in
the response of different bee species can produce varied responses to habitat
modification (Klein et al. 2003b; Cane et al. 2006; Greenleaf & Kremen 2006;
Winfree et al. 2007), and generalisations regarding effects of particular plant
reproductive characteristics on responses to land use changes are difficult (Aizen et
al. 2002; Ashworth et al. 2004). In particular, generalist honeybees may compensate
for reduced pollination by specialist native bees (Aizen & Feinsinger 1994a, b).
ii)
Plant-fungal: Generally altered mycorrhizal composition and functioning.
Changes in AMF composition and functioning due to management practices have
frequently been shown (Abbott & Robson 1991; Helgason et al. 1998; Borstler et al.
2006; Gosling et al. 2006; Hijri et al. 2006; Mathimaran et al. 2007; Stromberger et
al. 2007). Although AMF diversity has been shown to decline in modified habitats
(Opik et al. 2006), and fragmentation affects AMF community composition (Mangan
et al. 2004), no effects of patch size (Mangan et al. 2004) or edge effects (Mills 1995)
on AMF diversity have yet been shown. Diversity and composition of EMF
communities can be affected by both fragment size and isolation (Peay et al. 2007).
iii)
Seed dispersal: Reduction in dispersal. Habitat fragmentation has been
shown to lead to reduced bird frugivory (seed dispersal) (Valdivia & Simonetti 2007),
potentially causing a decline in obligately animal-dispersed trees (Laurance et al.
2006), and increased post-dispersal seed predation by rodents (Garcia & Chacoff
2007; but see Valdivia & Simonetti 2007). Although there are negative effects of
isolation, increased connectivity (corridors) between patches can facilitate seed
dispersal (Tewksbury et al. 2002). Habitat loss and fragmentation strongly affect large
species of mammals and birds that are highly mobile and responsible for the few
events of long-distance dispersal in fragmented habitats (Dirzo & Miranda 1990).
iv)
Plant-plant competition: Shift in competitive ability. Modification and
fragmentation of natural habitats has been shown frequently to alter local conditions
to favour competitive dominance of some plant species over others (Lavorel et al.
1997; Tilman & Lehman 2001; Dolt et al. 2005; Laurance et al. 2006; McEuen &
Curran 2006; Spiegelberger et al. 2006), and these effects may persist for decades
28
after management practices have ceased (Fraterrigo et al. 2006a, b). Increased grazing
intensity can also shift the interaction between plant species from competitive to
facilitative, when an unpalatable species provides protection to its neighbours (Graff
et al. 2007).
v)
Plant-hemiparasite: Possible benefit to parasite. Hemiparasites can benefit
from reduced competition following haymaking (Ameloot et al. 2006) or low levels
of simulated grazing (Hellstrom et al. 2004). More research on the effects of land use
changes on hemiparasites is required.
vi)
Plant-herbivore: General increase in herbivory, but variable. The inability
of gastropod herbivores to disperse from fragments has been shown to result in
increased herbivory (Stoll et al. 2006). Further studies have shown increased
herbivory (McEuen & Curran 2006) and herbivore abundance (Ryall & Fahrig 2005)
in more isolated fragments, and increased gall formation after a threshold decline in
forest cover (Chust et al. 2007). Land use intensity is also commonly associated with
increased herbivore abundance (Root 1973; Chen & Welter 2002; Klein et al. 2002;
Roschewitz et al. 2005). Although herbivores are less affected by patch size and
isolation than are higher trophic levels (Kruess & Tscharntke 1994; Tscharntke &
Brandl 2004; Valladares et al. 2006; Elzinga et al. 2007), they can still be negatively
affected. Loss of understorey forest habitat can lead to herbivore population declines
(Keeler et al. 2006), and grazing can affect herbivore community composition
(Hartley et al. 2003). Studies have also found lower aphid densities in simplified
landscapes (Roschewitz et al. 2005; Rand & Tscharntke 2007), and reduced
infestation rates by agromyzid flies on thistles in landscapes with reduced non-crop
area (Kruess 2003).
vii)
Plant-pathogen: Possible decrease, but variable, depending on scale of
observation. Pathogen occurrence may decline near fragment edges (Siitonen et al.
2005) or on small fragmented plant populations (Colling & Matthies 2004), but when
present, pathogen prevalence within the population can increase in fragmented
habitats (Groppe et al. 2001; Carlsson-Graner & Thrall 2002), possibly due to a
switch from the asymptomatic to the symptomatic state or to increased horizontal
pathogen transmission in fragments (Groppe et al. 2001). Increased connectivity
between sites created by the presence of roads has also been shown to increase
invasion success of a plant pathogen (Jules et al. 2002). Fragmentation is unlikely to
affect genetic susceptibility to pathogens (Galeuchet et al. 2005), but it can affect
abundance of vectors (Fabre et al. 2005; Grilli & Bruno 2007).
viii) Seed predation: Highly variable. Higher densities of rodents, subsidised by
agricultural habitats have been shown to provide increased levels of seed predation
(Jules & Rathcke 1999; Donoso et al. 2003; Tallmon et al. 2003; Garcia & Chacoff
2007). However, other studies have observed no direct effect of landscape
intensification on attack by weevils (Rand & Louda 2004), or even reduced seed
predation in fragmented habitats (Kruess & Tscharntke 1994; Steffan-Dewenter et al.
2001; Colling & Matthies 2004; Orrock & Damschen 2005; Ostergard & Ehrlen
2005), with negative effects being more likely for specialist seed-head feeders.
Grassland intensification can lead to reduced floral diversity, and subsequent
reductions in the quantity and diversity of grass and broad-leaved seed produced may
affect seed predators (Wilson et al. 1999).
29
ix)
Parasite-host: General increase in infection. Habitat modification is a
leading cause of the emergence of zoonoses (Chomel et al. 2007), and can promote
spread of vertebrate arboviruses (Toussaint et al. 2006). Although urbanization can in
some cases reduce the abundance of many wildlife parasites (Deplazes et al. 2004;
Fisher et al. 2005; Bradley & Altizer 2007), transmission can increase among hosts
adapted to urban (Bradley & Altizer 2007) or agricultural (Gilbert et al. 2007)
habitats. Therefore, parasite abundance, disease transmission and resulting population
decline are generally increased by urbanization (Prange et al. 2003; Riley et al. 2004;
Dhondt et al. 2005; Farnsworth et al. 2005; Wright & Gompper 2005; Ezenwa et al.
2006; Gibbs et al. 2006; Grieco et al. 2006; Yanoviak et al. 2006; Gilbert et al. 2007).
The abundance of wildlife parasites has also been shown to increase in fragmented
habitats (Allan et al. 2003; LoGiudice et al. 2003). Vectors that benefit from these
habitats can also spread diseases to rarer wildlife or to human populations (Grieco et
al. 2006; Yanoviak et al. 2006).
x)
Animal-animal competition: Shift in competitive ability. Habitat loss and
fragmentation can shift the competitive balance between different coccinelid (Zaviezo
et al. 2006), ant (Dauber & Wolters 2005), or generalist vs. specialist parasitoid
(Elzinga et al. 2007) species. In particular, differences in the colonisation ability of
different herbivore (Hines et al. 2005) and pollinator (Steffan-Dewenter et al. 2002)
species can affect their relative abundance in fragmented patches. Grazing intensity
can cause dominance shifts in herbivore communities (Hartley et al. 2003). Humaninduced desertification has been shown to shift the balance between specialist and
generalist lizards (Attum et al. 2006).
xi)
Predator-prey or parasitoid-host: Highly variable, depending on taxon.
Loss and fragmentation of natural habitats generally has its strongest negative effect
on specialists at higher trophic levels (such as insect parasitoids; Kruess & Tscharntke
1994; Bascompte & Solé 1998; Tscharntke & Brandl 2004; Valladares et al. 2006;
Elzinga et al. 2007). This can lead to reduced predator/prey ratios (Klein et al. 2002;
Hines et al. 2005; Ryall & Fahrig 2006; Watts & Didham 2006), rates of parasitism
(Roland & Taylor 1997; Thies & Tscharntke 1999; Kruess & Tscharntke 2000) and
ability of natural enemies to track increased density of their prey (Chen & Welter
2002). However, some studies have found no effects of fragmentation on parasitoidhost interactions (Chust et al. 2007), and the loss of specialist predators does not
necessarily imply an overall ecosystem-level reduction in predation (Swihart et al.
2001). In particular, landscape scale agricultural conversion frequently increases rates
of nest predation and brood parasitism (e.g., Andren 1992; Tewksbury et al. 2006),
particularly at edges. Similarly, parasitoid-host food web structure can be altered
significantly by increasing land use intensity, resulting in dominance of few
interactions, higher parasitoid/host ratios, and increased rates of parasitism of bees
and wasps (Tylianakis et al. 2007). These positive effects on predation rates may be
due to a subsidising effect of highly productive agricultural habitats on generalist
natural enemies (Rand et al. 2006), a shift in species composition to disturbed habitat
specialists (Andren 1992), or structural changes to the habitat affecting exposure of
prey to predation (Thompson & Gese 2007). Such positive effects of habitat
modification on predation by generalists may compensate or even overwhelm the
negative effects on specialist predators (Rand & Tscharntke 2007). Complex
multitrophic interactions can also blur the effects of fragmentation on specific
30
interactions. For example, landscape fragmentation was shown to reduce predation of
a woodpecker by facilitating predation of its mammalian predators by a goshawk
(Pakkala et al. 2006). The response of insect predator-prey interactions to land use
change may show significant variability depending on the type of study (experimental
vs. observational) and spatial scale (van Nouhuys 2005).
xii)
Decomposer food web: General reduction in biomass. Land use
intensification generally causes reductions of decomposer organisms, including both
microbial biomass (the basal consumer trophic level; Yeates et al. 1997; Frey et al.
1999; Wardle et al. 1999; Emmerling et al. 2001) and invertebrates occupying higher
trophic levels (Yeates et al. 1997; Wardle et al. 1999; Yeates et al. 1999; Doles et al.
2001; Schmidt et al. 2001; Cortet et al. 2002; Mulder et al. 2003; Wu et al. 2005; Adl
et al. 2006; Brennan et al. 2006; Chauvat et al. 2007), although a few studies have
reported no change in fungal (Elmholt & Labouriau 2005) or bacterial biomass (Frey
et al. 1999). This applies to both conversion of forest or grassland to agriculture
(Wardle 2002) and intensification of agricultural practice (e.g., through cultivation)
(Hendrix et al. 1986), and habitat restoration may not reverse these effects (Kardol et
al. 2005). Some components of the decomposer food web are far more adversely
affected than others, for example the fungal-based (vs. bacterial-based) energy
channel (Wardle 2002) and soil animals with larger body sizes (Wardle 1995).
Consequently, shifts in the fungal (Wu et al. 2007), nematode (Mulder et al. 2003)
and microflora (Bardgett et al. 2001) community structure have been recorded with
increased grazing intensity. The effects of other components of land use such as
habitat fragmentation remain largely unknown, although reductions in diversity of the
decomposer community (Rantalainen et al. 2005), abundance of microarthropods and
fungal biomass (Rantalainen et al. 2006) have been recorded in experimentally
fragmented habitats at small spatial scales. In contrast, other research has shown no
change in microarthropod density and diversity (Hoyle & Harborne 2005; Schneider
et al. 2007), and increases in soil predator density (Schneider et al. 2007) with habitat
isolation.
31
Figure S6: All drivers combined
32
References
Abbott L.K. & Robson A.D. (1991). Factors influencing the occurrence of vesicular arbuscular
mycorrhizas. Agric. Ecosyst. Environ., 35, 121-150.
Abrell L., Guerenstein P.G., Mechaber W.L., Stange G., Christensen T.A., Nakanishi K. & Hildebrand
J.G. (2005). Effect of elevated atmospheric CO2 on oviposition behavior in Manduca sexta
moths. Glob. Change Biol., 11, 1272-1282.
Adl S.M., Coleman D.C. & Read F. (2006). Slow recovery of soil biodiversity in sandy loam soils of
Georgia after 25 years of no-tillage management. Agric. Ecosyst. Environ., 114, 323-334.
Aflakpui G.K.S., Gregory P. & Froud-Williams R.J. (2005). Carbon (C-13) and nitrogen (N-15)
translocation in a maize-Striga hermonthica association. Exp. Agric., 41, 321-333.
Aguilar R., Ashworth L., Galetto L. & Aizen M.A. (2006). Plant reproductive susceptibility to habitat
fragmentation: review and synthesis through a meta-analysis. Ecol. Lett., 9, 968-980.
Aizen M.A., Ashworth L. & Galetto L. (2002). Reproductive success in fragmented habitats: do
compatibility systems and pollination specialization matter? J. Vege. Sci., 13, 885-892.
Aizen M.A. & Feinsinger P. (1994a). Forest fragmentation, pollination, and plant reproduction in
Chaco dry forest, Argentina. Ecology, 75, 330-51.
Aizen M.A. & Feinsinger P. (1994b). Habitat fragmentation, native insect pollinators, and feral honey
bees in Argentine "Chaco Serrano". Ecol. Appl., 4, 378-92.
Alberton O., Kuyper T.W. & Gorissen A. (2005). Taking mycocentrism seriously: mycorrhizal fungal
and plant responses to elevated CO2. New Phytol., 167, 859-868.
Aldea M., Hamilton J.G., Resti J.P., Zangerl A.R., Berenbaum M.R., Frank T.D. & DeLucia E.H.
(2006). Comparison of photosynthetic damage from arthropod herbivory and pathogen
infection in understory hardwood saplings. Oecologia, 149, 221-232.
Allan B.F., Keesing F. & Ostfeld R.S. (2003). Effect of forest fragmentation on Lyme disease risk.
Cons. Biol., 17, 267-272.
Allen M.F., Klironomos J.N., Treseder K.K. & Oechel W.C. (2005). Responses of soil biota to elevated
CO2 in a chaparral ecosystem. Ecol. Appl., 15, 1701-1711.
Alyokhin A. & Sewell G. (2004). Changes in a lady beetle community following the establishment of
three alien species. Biol. Inv., 6, 463-471.
Ameloot E., Verheyen K., Bakker J.P., De Vries Y. & Hermy M. (2006). Long-term dynamics of the
hemiparasite Rhinanthus angustifolius and its relationship with vegetation structure. J. Vege.
Sci., 17, 637-646.
Andren H. (1992). Corvid density and nest predation in relation to forest fragmentation - a landscape
perspective. Ecology, 73, 794-804.
Andrew N.R. & Hughes L. (2005). Herbivore damage along a latitudinal gradient: relative impacts of
different feeding guilds. Oikos, 108, 176-182.
Armolaitis K. (1998). Nitrogen pollution on the local scale in Lithuania: vitality of forest ecosystems.
Environ. Pollution, 102, 55-60.
Arnold S.S., Fernandez I.J., Rustad L.E. & Zibilske L.M. (1999). Microbial response of an acid forest
soil to experimental soil warming. Biol. Fert. Soils, 30, 239-244.
Ashworth L., Aguilar R., Galetto L. & Aizen M.A. (2004). Why do pollination generalist and specialist
plant species show similar reproductive susceptibility to habitat fragmentation? J. Ecol., 92,
717-719.
Asshoff R. & Hättenschwiler S. (2005). Growth and reproduction of the alpine grasshopper Miramella
alpina feeding on CO2-enriched dwarf shrubs at treeline. Oecologia, 142, 191-201.
Atkin O.K., Schortemeyer M., McFarlane N. & Evans J.R. (1999). The response of fast- and slowgrowing Acacia species to elevated atmospheric CO2: an analysis of the underlying
components of relative growth rate. Oecologia, 120, 544-554.
Attum O., Eason P., Cobbs G. & El Din S.M.B. (2006). Response of a desert lizard community to
habitat degradation: Do ideas about habitat specialists/generalists hold? Biol. Cons., 133, 5262.
Avis P.G. & Charvat I. (2005). The response of ectomycorrhizal fungal inoculum to long-term
increases in nitrogen supply. Mycologia, 97, 329-337.
Awmack C.S., Harrington R. & Lindroth R.L. (2004). Aphid individual performance may not predict
population responses to elevated CO2 or O-3. Glob. Change Biol., 10, 1414-1423.
Ayongwa G.C., Stomph T.J., Emechebe A.M. & Kuyper T.W. (2006). Root nitrogen concentration of
sorghum above 2% produces least Striga hermonthica seed stimulation. Ann. Appl. Biol., 149,
255-262.
33
Badgery W.B., Kemp D.R., Michalk D.L. & King W. (2005). Competition for nitrogen between
Australian native grasses and the introduced weed Nassella trichotoma. Ann. Bot., 96, 799809.
Bale J.S., Masters G.J., Hodkinson I.D., Awmack C., Bezemer T.M., Brown V.K., Butterfield J., Buse
A., Coulson J.C., Farrar J., Good J.E.G., Harrington R., Hartley S., Jones T.H., Lindroth R.L.,
Press M.C., Symrnioudis I., Watt A.D. & Whittaker J.B. (2002). Herbivory in global climate
change research: direct effects of rising temperature on insect herbivores. Glob. Change Biol.,
8, 1-16.
Barbehenn R.V., Karowe D.N. & Spickard A. (2004). Effects of elevated atmospheric CO2 on the
nutritional ecology of C-3 and C-4 grass-feeding caterpillars. Oecologia, 140, 86-95.
Bardgett R.D., Jones A.C., Jones D.L., Kemmitt S.J., Cook R. & Hobbs P.J. (2001). Soil microbial
community patterns related to the history and intensity of grazing in sub-montane ecosystems.
Soil Biol. Biochem., 33, 1653-1664.
Bardgett R.D., Kandeler E., Tscherko D., Hobbs P.J., Bezemer T.M., Jones T.H. & Thompson L.J.
(1999a). Below-ground microbial community development in a high temperature world.
Oikos, 85, 193-203.
Bardgett R.D., Mawdsley J.L., Edwards S., Hobbs P.J., Rodwell J.S. & Davies W.J. (1999b). Plant
species and nitrogen effects on soil biological properties of temperate upland grasslands.
Funct. Ecol., 13, 650-660.
Baruch Z. & Jackson R.B. (2005). Responses of tropical native and invader C-4 grasses to water stress,
clipping and increased atmospheric CO2 concentration. Oecologia, 145, 522-532.
Bascompte J. & Solé R.V. (1998). Effects of habitat destruction in a prey-predator metapopulation
model. J. Theor. Biol., 195, 383-393.
Batten K.M., Scow K.M., Davies K.F. & Harrison S.P. (2006). Two invasive plants alter soil microbial
community composition in serpentine grasslands. Biol. Inv., 8, 217-230.
Battisti A., Stastny M., Netherer S., Robinet C., Schopf A., Roques A. & Larsson S. (2005). Expansion
of geographic range in the pine processionary moth caused by increased winter temperatures.
Ecol. Appl., 15, 2084-2096.
Baum C. & Makeschin F. (2000). Effects of nitrogen and phosphorus fertilization on mycorrhizal
formation of two poplar clones (Populus trichocarpa and P. tremula x tremuloides). J. Plant
Nutr. Soil Sci.-Z. Pflanzenernahr. Bodenkd., 163, 491-497.
Baum C., Weih M., Verwijst T. & Makeschin F. (2002). The effects of nitrogen fertilization and soil
properties on mycorrhizal formation of Salix viminalis. Forest Ecol. Manag., 160, 35-43.
Bazzaz F.A. (1990). The response of natural ecosystems to the rising global CO2 levels. Ann. Rev.
Ecol. Syst., 21, 167-196.
Belnap J., Phillips S.L., Sherrod S.K. & Moldenke A. (2005). Soil biota can change after exotic plant
invasion: does this affect ecosystem processes? Ecology, 86, 3007-3017.
Benson J., Van Driesche R.G., Pasquale A. & Elkinton J. (2003). Introduced braconid parasitoids and
range reduction of a native butterfly in New England. Biol. Control, 28, 197-213.
Berntson G.M., Rajakaruna N. & Bazzaz F.A. (1998). Growth and nitrogen uptake in an experimental
community of annuals exposed to elevated atmospheric CO2. Glob. Change Biol., 4, 607-626.
Bezemer T.M. & Jones T.H. (1998). Plant-insect herbivore interactions in elevated atmospheric CO2:
quantitative analyses and guild effects. Oikos, 82, 212-222.
Bezemer T.M., Jones T.H. & Knight K.J. (1998). Long-term effects of elevated CO2 and temperature
on populations of the peach potato aphid Myzus persicae and its parasitoid Aphidius
matricariae. Oecologia, 116, 128-135.
Bezemer T.M., Knight K.J., Newington J.E. & Jones T.H. (1999). How general are aphid responses to
elevated atmospheric CO2? Ann. Ent. Soc. Am., 92, 724-730.
Bidart-Bouzat M.G. (2004). Herbivory modifies the lifetime fitness response of Arabidopsis thaliana to
elevated CO2. Ecology, 85, 297-303.
Bigger D.S. (1999). Consequences of patch size and isolation for a rare plant: Pollen limitation and
seed predation. Nat. Areas J., 19, 239-244.
Blancafort X. & Gomez C. (2005). Consequences of the Argentine ant, Linepithema humile (Mayr),
invasion on pollination of Euphorbia characias (L.) (Euphorbiaceae). Acta Oecol., 28, 49-55.
Blanche R. & Cunningham S.A. (2005). Rain forest provides pollinating beetles for atemoya crops. J.
Econ. Ent., 98, 1193-1201.
Boag B. (2000). The impact of the New Zealand flatworm on earthworms and moles in agricultural
land in western Scotland. Aspects Appl. Biol., 62, 79-84.
Bobbink R., Hornung M. & Roelofs J.G.M. (1998). The effects of air-borne nitrogen pollutants on
species diversity in natural and semi-natural European vegetation. J. Ecol., 86, 717-738.
34
Bohlen P.J., Scheu S., Hale C.M., McLean M.A., Migge S., Groffman P.M. & Parkinson D. (2004).
Non-native invasive earthworms as agents of change in northern temperate forests. Front.
Ecol. Environ., 2, 427-435.
Borstler B., Renker C., Kahmen A. & Buscot F. (2006). Species composition of arbuscular mycorrhizal
fungi in two mountain meadows with differing management types and levels of plant
biodiversity. Biol. Fert. Soils, 42, 286-298.
Bosch M., Simon J., Rovira A.M., Molero J. & Blanche C. (2002). Pollination ecology of the prePyrenean endemic Petrocoptis montsicciana (Caryophyllaceae): effects of population size.
Biol. J. Linn. Soc., 76, 79-90.
Both C., Bouwhuis S., Lessells C.M. & Visser M.E. (2006). Climate change and population declines in
a long-distance migratory bird. Nature, 441, 81-83.
Bradley C.A. & Altizer S. (2007). Urbanization and the ecology of wildlife diseases. Trends Ecol.
Evol., 22, 95-102.
Bradley K., Drijber R.A. & Knops J. (2006). Increased N availability in grassland soils modifies their
microbial communities and decreases the abundance of arbuscular mycorrhizal fungi. Soil
Biol. Biochem., 38, 1583-1595.
Brennan A., Fortune T. & Bolger T. (2006). Collembola abundances and assemblage structures in
conventionally tilled and conservation tillage arable systems. Pedobiologia, 50, 135-145.
Briones M.J.I., Ineson P. & Piearce T.G. (1997). Effects of climate change on soil fauna; responses of
enchytraeids, Diptera larvae and tardigrades in a transplant experiment. Appl. Soil Ecol., 6,
117-134.
Brooker R.W. (2006). Plant-plant interactions and environmental change. New Phytol., 171, 271-284.
Brooks M.L. (2003). Effects of increased soil nitrogen on the dominance of alien annual plants in the
Mojave Desert. J. Appl. Ecol., 40, 344-353.
Brown B.J., Mitchell R.J. & Graham S.A. (2002). Competition for pollination between an invasive
species (purple loosestrife) and a native congener. Ecology, 83, 2328-2336.
Bruna E.M. & Kress W.J. (2002). Habitat fragmentation and the demographic structure of an
Amazonian understory herb (Heliconia acuminata). Cons. Biol., 16, 1256-1266.
Brys R., Jacquemyn H., Endels P., van Rossum F., Hermy M., Triest L., De Bruyn L. & Blust G.D.E.
(2004). Reduced reproductive success in small populations of the self-incompatible Primula
vulgaris. J. Ecol., 92, 5-14.
Bunemann E.K., Schwenke G.D. & Van Zwieten L. (2006). Impact of agricultural inputs on soil
organisms - a review. Aust. J. Soil Res., 44, 379-406.
Burtelow A.E., Bohlen P.J. & Groffman P.M. (1998). Influence of exotic earthworm invasion on soil
organic matter, microbial biomass and denitrification potential in forest soils of the
northeastern United States. Appl. Soil Ecol., 9, 197-202.
Byers J.E. & Noonburg E.G. (2003). Scale dependent effects of biotic resistance to biological invasion.
Ecology, 84, 1428-1433.
Callaway R.M., DeLuca T.H. & Belliveau W.M. (1999). Biological-control herbivores may increase
competitive ability of the noxious weed Centaurea maculosa. Ecology, 80, 1196-1201.
Cane J.H., Minckley R.L., Kervin L.J., Roulston T.H. & Williams N.M. (2006). Complex responses
within a desert bee guild (Hymenoptera: Apiformes) to urban habitat fragmentation. Ecol.
Appl., 16, 632-644.
Carlsson-Graner U. & Thrall P.H. (2002). The spatial distribution of plant populations, disease
dynamics and evolution of resistance. Oikos, 97, 97-110.
Carney S.E., Byerley M.B. & Holway D.A. (2003). Invasive Argentine ants (Linepithema humile) do
not replace native ants as seed dispersers of Dendromecon rigida (Papaveraceae) in
California, USA. Oecologia, 135, 576-582.
Cattadori I.M., Haydon D.T. & Hudson P.J. (2005). Parasites and climate synchronize red grouse
populations. Nature, 433, 737-741.
Celebrezze T. & Paton D.C. (2004). Do introduced honeybees (Apis mellifera, Hymenoptera) provide
full pollination service to bird-adapted Australian plants with small flowers? An experimental
study of Brachyloma ericoides (Epacridaceae). Austral Ecol., 29, 129-136.
Chacoff N.P. & Aizen M.A. (2006). Edge effects on flower-visiting insects in grapefruit plantations
bordering premontane subtropical forest. J. Appl. Ecol., 43, 18-27.
Chacoff N.P., Aizen M.A. & Aschero V. (2008). Proximity to forest edge does not affect crop
production despite pollen limitation. Proc. R. Soc. Lond. B, 275, 907-913.
Chakraborty S., Tiedemann A.V. & Teng P.S. (2000). Climate change: potential impact on plant
diseases. Environ. Pollution, 108, 317-326.
35
Chauvat M., Wolters V. & Dauber J. (2007). Response of collembolan communities to land-use change
and grassland succession. Ecography, 30, 183-192.
Chauvel A., Grimaldi M., Barros E., Blanchart E., Desjardins T., Sarrazin M. & Lavelle P. (1999).
Pasture damage by an Amazonian earthworm. Nature, 398, 32-33.
Chen F.J., Ge F. & Parajulee M.N. (2005). Impact of elevated CO2 on tri-trophic interaction of
Gossypium hirsutum, Aphis gossypii, and Leis axyridis. Environ. Ent., 34, 37-46.
Chen Y.H. & Welter S.C. (2002). Abundance of a native moth Homoeosoma electellum (Lepidopter :
Pyralidae) and activity of indigenous parasitoids in native and agricultural sunflower habitats.
Environ. Ent., 31, 626-636.
Cheptou P.O. & Avendano L.G. (2006). Pollination processes and the Allee effect in highly fragmented
populations: consequences for the mating system in urban environments. New Phytol., 172,
774-783.
Chittka L. & Schurkens S. (2001). Successful invasion of a floral market - An exotic Asian plant has
moved in on Europe's river-banks by bribing pollinators. Nature, 411, 653-653.
Chomel B.B., Belotto A. & Meslin F.X. (2007). Wildlife, exotic pets, and emerging zoonoses. Emerg.
Infec. Diseas., 13, 6-11.
Chong J.H. & Oetting R.D. (2006). Influence of temperature, nourishment, and storage period on the
longevity and fecundity of the mealybug parasitoid, Anagyrus sp. nov. nr. sinope Noyes and
Menezes (Hymenoptera : Encyrtidae). Environ. Ent., 35, 1198-1207.
Chong J.H., Oetting R.D. & Osborne L.S. (2005). Development of Diomus austrinus Gordon
(Coleoptera : Coccinellidae) on two mealybug prey species at five constant temperatures. Biol.
Control, 33, 39-48.
Chong J.H., van Iersel M.W. & Oetting R.D. (2004). Effects of elevated carbon dioxide levels and
temperature on the life history of the Madeira mealybug (Hemiptera : Pseudococcidae). J. Ent.
Sci., 39, 387-397.
Christian C.E. (2001). Consequences of a biological invasion reveal the importance of mutualism for
plant communities. Nature, 413, 635-639.
Chung H.G., Zak D.R. & Lilleskov E.A. (2006). Fungal community composition and metabolism under
elevated CO2 and O-3. Oecologia, 147, 143-154.
Chust G., Garbin L. & Pujade-Villar J. (2007). Gall wasps and their parasitoids in cork oak fragmented
forests. Ecol. Ent., 32, 82-91.
Cipollini M.L., Paulk E., Mink K., Vaughn K. & Fischer T. (2004). Defense tradeoffs in fleshy fruits:
Effects of resource variation on growth, reproduction, and fruit secondary chemistry in
Solanum carolinense. J. Chem. Ecol., 30, 1-17.
Clark H., Newton P.C.D., Bell C.C. & Glasgow E.M. (1997). Dry matter yield, leaf growth and
population dynamics in Lolium perenne, Trifolium repens-dominated pasture turves exposed
to two levels of elevated CO2. J. Appl. Ecol., 34, 304-316.
Cleland E.E., Peters H.A., Mooney H.A. & Field C.B. (2006). Gastropod herbivory in response to
elevated CO2 and N addition impacts plant community composition. Ecology, 87, 686-694.
Clemmensen K.E., Michelsen A., Jonasson S. & Shaver G.R. (2006). Increased ectomycorrhizal fungal
abundance after long-term fertilization and warming of two arctic tundra ecosystems. New
Phytol., 171, 391-404.
Cole L., Bardgett R.D., Ineson P. & Adamson J.K. (2002). Relationships between enchytraeid worms
(Oligochaeta), climate change, and the release of dissolved organic carbon from blanket peat
in northern England. Soil Biol. Biochem., 34, 599-607.
Colling G. & Matthies D. (2004). The effects of plant population size on the interactions between the
endangered plant Scorzonera humilis, a specialised herbivore, and a phytopathogenic fungus.
Oikos, 105, 71-78.
Convey P., Pugh P.J.A., Jackson C., Murray A.W., Ruhland C.T., Xiong F.S. & Day T.A. (2002).
Response of antarctic terrestrial microarthropods to long-term climate manipulations. Ecology,
83, 3130-3140.
Cornelissen T. & Stiling P. (2006a). Does low nutritional quality act as a plant defence? An
experimental test of the slow-growth, high-mortality hypothesis. Ecol. Ent., 31, 32-40.
Cornelissen T. & Stiling P. (2006b). Responses of different herbivore guilds to nutrient addition and
natural enemy exclusion. EcoSci., 13, 66-74.
Cortet J., Ronce D., Poinsot-Balaguer N., Beaufreton C., Chabert A., Viaux P. & de Fonseca J.P.C.
(2002). Impacts of different agricultural practices on the biodiversity of microarthropod
communities in arable crop systems. Eur. J. Soil Biol., 38, 239-244.
Courchamp F., Woodroffe R. & Roemer G.W. (2003). Removing protected populations to save
endangered species. Science, 302, 1532.
36
Cross J.R. (1981). The establishment of Rhododendron ponticum in the Killarney Oakwoods, SW
Ireland. J. Ecol., 69, 807-824.
Dag A. & Eisikowitch D. (2000). The effect of carbon dioxide enrichment on nectar production in
melons under greenhouse conditions. J. Apic. Res., 39, 88-89.
Dale H. & Press M.C. (1998). Elevated atmospheric CO2 influences the interaction between the
parasitic angiosperm Orobanche minor and its host Trifolium repens. New Phytol., 140, 65-73.
Darvill B., Ellis J.S., Lye G.C. & Goulson D. (2006). Population structure and inbreeding in a rare and
declining bumblebee, Bombus muscorum (Hymenoptera: Apidae). Molec. Ecol., 15, 601-611.
Dauber J. & Wolters V. (2005). Colonization of temperate grassland by ants. Basic Appl. Ecol., 6, 8391.
Davis A.R. (2003a). Influence of elevated CO2 and ultraviolet-B radiation levels on floral nectar
production: a nectary-morphological perspective. Plant Syst. Evol., 238, 169-181.
Davis M.A. (2003b). Biotic globalization: does competition from introduced species threaten
biodiversity? BioScience, 53, 481-489.
de Vries F.T., Hoffland E., van Eekeren N., Brussaard L. & Bloem J. (2006). Fungal/bacterial ratios in
grasslands with contrasting nitrogen management. Soil Biol. Biochem., 38, 2092-2103.
Deplazes P., Hegglin D., Gloor S. & Romig T. (2004). Wilderness in the city: the urbanization of
Echinococcus multilocularis. Trends Parasitol., 20, 77-84.
DeVault T.L., Brisbin I.L. & Rhodes O.E. (2004). Factors influencing the acquisition of rodent carrion
by vertebrate scavengers and decomposers. Can. J. Zool., 82, 502-509.
Dhondt A.A., Altizer S., Cooch E.G., Davis A.K., Dobson A., Driscoll M.J.L., Hartup B.K., Hawley
D.M., Hochachka W.M., Hosseini P.R., Jennelle C.S., Kollias G.V., Ley D.H., Swarthout
E.C.H. & Sydenstricker K.V. (2005). Dynamics of a novel pathogen in an avian host:
Mycoplasmal conjunctivitis in house finches. Acta Trop., 94, 77-93.
Diaz S., Fraser L.H., Grime J.P. & Falczuk V. (1998). The impact of elevated CO2 on plant-herbivore
interactions: experimental evidence of moderating effects at the community level. Oecologia,
117, 177-186.
Diekotter T., Haynes K.J., Mazeffa D. & Crist T.O. (2007). Direct and indirect effects of habitat area
and matrix composition on species interactions among flower-visiting insects. Oikos, 116,
1588-1598.
Diez J. (2005). Invasion biology of Australian ectomycorrhizal fungi introduced with eucalypt
plantations into the Iberian Peninsula. Biol. Inv., 7, 3-15.
Dijkstra F.A., Hobbie S.E., Reich P.B. & Knops J.M.H. (2005). Divergent effects of elevated CO2, N
fertilization, and plant diversity on soil C and N dynamics in a grassland field experiment.
Plant Soil, 272, 41-52.
Dirzo R. & Miranda A. (1990). Contemporary neotropical defaunation and forest structure, function,
and diversity - a sequel. Cons. Biol., 4, 444-447.
Docherty M., Wade F.A., Hurst D.K., Whittaker J.B. & Lea P.J. (1997). Responses of tree sap-feeding
herbivores to elevated CO2. Glob. Change Biol., 3, 51-59.
Doles J.L., Zimmerman R.J. & Moore J.C. (2001). Soil microarthropod community structure and
dynamics in organic and conventionally managed apple orchards in Western Colorado, USA.
Appl.Soil Ecol., 18, 83-96.
Dollery R., Hodkinson I.D. & Jonsdottir I.S. (2006). Impact of warming and timing of snow melt on
soil microarthropod assemblages associated with Dryas-dominated plant communities on
Svalbard. Ecography, 29, 111-119.
Dolt C., Goverde M. & Baur B. (2005). Effects of experimental small-scale habitat fragmentation on
above-and below-ground plant biomass in calcareous grasslands. Acta Oecol., 27, 49-56.
Donoso D.S., Grez A.A. & Simonetti J.A. (2003). Effects of forest fragmentation on the granivory of
differently sized seeds. Biol. Cons., 115, 63-70.
Duncan D.H., Nicotra A.B., Wood J.T. & Cunningham S.A. (2004). Plant isolation reduces outcross
pollen receipt in a partially self-compatible herb. J. Ecol., 92, 977-985.
Dunnett N.P. & Grime J.P. (1999). Competition as an amplifier of short-term vegetation responses to
climate: an experimental test. Funct. Ecol., 13, 388-395.
Durant J.M., Anker-Nilssen T. & Stenseth N.C. (2003). Trophic interactions under climate fluctuations:
the Atlantic puffin as an example. Proc. R. Soc. Lond. B, 270, 1461-1466.
Durant J.M., Hjermann D.O., Anker-Nilssen T., Beaugrand G., Mysterud A., Pettorelli N. & Stenseth
N.C. (2005). Timing and abundance as key mechanisms affecting trophic interactions in
variable environments. Ecol. Lett., 8, 952-958.
37
Dwyer G.I., Gibbs M.J., Gibbs A.J. & Jones R.A.C. (2007). Wheat streak mosaic virus in Australia:
Relationship to isolates from the Pacific Northwest of the USA and its dispersion via seed
transmission. Plant Dis., 91, 164-170.
Ebi K.L., Hartman J., Chan N., McConnell J., Schlesinger M. & Weyant J. (2005). Climate suitability
for stable malaria transmission in Zimbabwe under different climate change scenarios. Clim.
Change, 73, 375-393.
Edwards G.R., Bourdot G.W. & Crawley M.J. (2000). Influence of herbivory, competition and soil
fertility on the abundance of Cirsium arvense in acid grassland. J. Appl. Ecol., 37, 321-334.
Ehrenfeld J.G. (2003). Effects of exotic plant invasions on soil nutrient cycling processes. Ecosyst., 6,
503-523.
El-Hajj Z., Kavanagh K., Rose C. & Kanaan-Atallah Z. (2004). Nitrogen and carbon dynamics of a
foliar biotrophic fungal parasite in fertilized Douglas-fir. New Phytol., 163, 139-147.
Elkinton J.S., Parry D. & Boettner G.H. (2006). Implicating an introduced generalist parasitoid in the
invasive browntail moth's enigmatic demise. Ecology, 87, 2664-2672.
Ellis J.S., Knight M.E., Darvill B. & Goulson D. (2006). Extremely low effective population sizes,
genetic structuring and reduced genetic diversity in a threatened bumblebee species, Bombus
sylvarum (Hymenoptera : Apidae). Molec. Ecol., 15, 4375-4386.
Elmholt S. & Labouriau R. (2005). Fungi in Danish soils under organic and conventional farming.
Agric. Ecosyst. Environ., 107, 65-73.
Elzinga J.A., van Nouhuys S., van Leeuwen D.J. & Biere A. (2007). Distribution and colonisation
ability of three parasitoids and their herbivorous host in a fragmented landscape. Basic Appl.
Ecol., 8, 75-88.
Emmerling C., Udelhoven T. & Schroder D. (2001). Response of soil microbial biomass and activity to
agricultural de-intensification over a 10 year period. Soil Biol. Biochem., 33, 2105-2114.
Erelli M.C., Ayres M.P. & Eaton G.K. (1998). Altitudinal patterns in host suitability for forest insects.
Oecologia, 117, 133-142.
Erhardt A., Rusterholz H.P. & Stocklin J. (2005). Elevated carbon dioxide increases nectar production
in Epilobium angustifolium L. Oecologia, 146, 311-317.
Ettema C.H., Lowrance R. & Coleman D.C. (1999). Riparian soil response to surface nitrogen input:
the indicator potential of free-living soil nematode populations. Soil Biol. Biochem., 31, 16251638.
Evans E.W. (2004). Habitat displacement of North American ladybirds by an introduced species.
Ecology, 85, 637-647.
Evans E.W. & England S. (1996). Indirect interactions in biological control of insects: Pests and
natural enemies in alfalfa. Ecol. Appl., 6, 920-930.
Ezenwa V.O., Godsey M.S., King R.J. & Guptill S.C. (2006). Avian diversity and West Nile virus:
testing associations between biodiversity and infectious disease risk. Proc. R. Soc. Lond. B,
273, 109-117.
Fabre F., Plantegenest M., Mieuzet L., Dedryver C.A., Leterrier J.L. & Jacquot E. (2005). Effects of
climate and land use on the occurrence of viruliferous aphids and the epidemiology of barley
yellow dwarf disease. Agric. Ecosyst. Environ., 106, 49-55.
Farnsworth M.L., Wolfe L.L., Hobbs N.T., Burnham K.P., Williams E.S., Theobald D.M., Conner
M.M. & Miller M.W. (2005). Human land use influences chronic wasting disease prevalence
in mule deer. Ecol. Appl., 15, 119-126.
Fellowes M.D.E., Kraaijeveld A.R. & Godfray H.C.J. (1999). Cross-resistance following artificial
selection for increased defense against parasitoids in Drosophila melanogaster. Evolution, 53,
966-972.
Ferguson R.N. & Drake D.R. (1999). Influence of vegetation structure on spatial patterns of seed
deposition by birds. N. Z. J. Bot., 37, 671-677.
Fisher C., Reperant L.A., Weber J.M., Hegglin D. & Deplazes P. (2005). Echinococcus multilocularis
infections of rural, residential and urban foxes (Vulpes vulpes) in the canton of Geneva,
Switzerland. Parasite-J. Soc. Fr. Parasitol., 12, 339-346.
Fisk M.C. & Fahey T.J. (2001). Microbial biomass and nitrogen cycling responses to fertilization and
litter removal in young northern hardwood forests. Biogeochem., 53, 201-223.
Fitter A.H. & Fitter R.S.R. (2002). Rapid changes in flowering time in British plants. Science, 296,
1689-1691.
Fluckiger W. & Braun S. (1998). Nitrogen deposition in Swiss forests and its possible relevance for
leaf nutrient status, parasite attacks and soil acidification. Environ. Pollution, 102, 69-76.
Folgarait P.J. & Sala O.E. (2002). Granivory rates by rodents, insects, and birds at different microsites
in the Patagonian steppe. Ecography, 25, 417-427.
38
Franks S.J., Kral A.M. & Pratt P.D. (2006). Herbivory by introduced insects reduces growth and
survival of Melaleuca quinquenervia seedlings. Environ. Ent., 35, 366-372.
Fransson P.M.A., Taylor A.F.S. & Finlay R.D. (2001). Elevated atmospheric CO2 alters root symbiont
community structure in forest trees. New Phytol., 152, 431-442.
Fraterrigo J.M., Turner M.G. & Pearson S.M. (2006a). Interactions between past land use, life-history
traits and understory spatial heterogeneity. Landscape Ecol., 21, 777-790.
Fraterrigo J.M., Turner M.G. & Pearson S.M. (2006b). Previous land use alters plant allocation and
growth in forest herbs. J. Ecol., 94, 548-557.
Frey S.D., Elliott E.T. & Paustian K. (1999). Bacterial and fungal abundance and biomass in
conventional and no-tillage agroecosystems along two climatic gradients. Soil Biol. Biochem.,
31, 573-585.
Fuhrer J. (2003). Agroecosystern responses to combinations of elevated CO 2, ozone, and global climate
change. Agric. Ecosyst. Environ., 97, 1-20.
Fukami T., Wardle D.A., Bellingham P.J., Mulder C.P.H., Towns D.R., Yeates G.W., Bonner K.I.,
Durrett M.S., Grant-Hoffman M.N. & Williamson W.M. (2006). Above- and below-ground
impacts of introduced predators in seabird-dominated island ecosystems. Ecol. Lett., 9, 12991307.
Gabriel D. & Tscharntke T. (2007). Insect pollinated plants benefit from organic farming. Agric.
Ecosyst. Environ., 118, 43-48.
Galeuchet D.J., Perret C. & Fischer M. (2005). Performance of Lychnis flos-cuculi from fragmented
populations under experimental biotic interactions. Ecology, 86, 1002-1011.
Gamper H., Hartwig U.A. & Leuchtmann A. (2005). Mycorrhizas improve nitrogen nutrition of
Trifolium repens after 8 yr of selection under elevated atmospheric CO2 partial pressure. New
Phytol., 167, 531-542.
Gamper H., Peter M., Jansa J., Luscher A., Hartwig U.A. & Leuchtmann A. (2004). Arbuscular
mycorrhizal fungi benefit from 7 years of free air CO 2 enrichment in well-fertilized grass and
legume monocultures. Glob. Change Biol., 10, 189-199.
Garcia D. & Chacoff N.P. (2007). Scale-dependent effects of habitat fragmentation on hawthorn
pollination. Cons. Biol., 21, 400-407.
Gardener M.C. & Gillman M.P. (2001). The effects of soil fertilizer on amino acids in the floral nectar
of corncockle, Agrostemma githago (Caryophyllaceae). Oikos, 92, 101-106.
Garrett K.A., Dendy S.P., Frank E.E., Rouse M.N. & Travers S.E. (2006). Climate change effects on
plant disease: Genomes to ecosystems. Annu. Rev. Phytopathol., 44, 489-509.
Gathmann A. & Tscharntke T. (2002). Foraging ranges of solitary bees. J. Anim. Ecol., 71, 757-764.
Gavito M.E., Curtis P.S., Mikkelsen T.N. & Jakobsen I. (2000). Atmospheric CO2 and mycorrhiza
effects on biomass allocation and nutrient uptake of nodulated pea (Pisum sativum L.) plants.
J. Exp. Bot., 51, 1931-1938.
Gavito M.E., Schweiger P. & Jakobsen I. (2003). P uptake by arbuscular mycorrhizal hyphae: effect of
soil temperature and atmospheric CO2 enrichment. Glob. Change Biol., 9, 106-116.
Geiger W., Alcorlo P., Baltanas A. & Montes C. (2005). Impact of an introduced Crustacean on the
trophic webs of Mediterranean wetlands. Biol. Inv., 7, 49-73.
Ghazoul J. (2004). Alien abduction: disruption of native plant-pollinator interactions by invasive
species. Biotropica, 36, 156-164.
Ghazoul J. (2005). Pollen and seed dispersal among dispersed plants. Biol. Rev., 80, 413-443.
Ghazoul J. & McLeish M. (2001). Reproductive ecology of tropical forest trees in logged and
fragmented habitats in Thailand and Costa Rica. Plant Ecol., 153, 335-345.
Gibbs S.E.J., Wimberly M.C., Madden M., Masour J., Yabsley M.J. & Stallknecht D.E. (2006). Factors
affecting the geographic distribution of West Nile virus in Georgia, USA: 2002-2004. VectorBorne Zoonotic Dis., 6, 73-82.
Gigord L., Picot F. & Shykoff J.A. (1999). Effects of habitat fragmentation on Dombeya acutangula
(Sterculiaceae), a native tree on La Réunion (Indian Ocean). Biol. Cons., 88, 43-51.
Gilbert M., Xiao X.M., Chaitaweesub P., Kalpravidh W., Premashthira S., Boles S. & Slingenbergh J.
(2007). Avian influenza, domestic ducks and rice agriculture in Thailand. Agric. Ecosyst.
Environ., 119, 409-415.
Gillespie I.G. & Allen E.B. (2006). Effects of soil and mycorrhizae from native and invaded vegetation
on a rare California forb. Appl. Soil Ecol., 32, 6-12.
Gosling P., Hodge A., Goodlass G. & Bending G.D. (2006). Arbuscular mycorrhizal fungi and organic
farming. Agric. Ecosyst. Environ., 113, 17-35.
Gotelli N.J. & Arnett A.E. (2000). Biogeographic effects of red fire ant invasion. Ecol. Lett., 3, 257261.
39
Graff P., Aguiar M.R. & Chaneton E.J. (2007). Shifts in positive and negative plant interactions along a
grazing intensity gradient. Ecology, 88, 188-199.
Gratton C. & Denno R.F. (2006). Arthropod food web restoration following removal of an invasive
wetland plant. Ecol. Appl., 16, 622-631.
Greenleaf S.S. & Kremen C. (2006). Wild bee species increase tomato production and respond
differently to surrounding land use in Northern California. Biol. Cons., 133, 81-87.
Greer D.H., Laing W.A., Campbell B.D. & Halligan E.A. (2000). The effect of perturbations in
temperature and photon flux density on the growth and photosynthetic responses of five
pasture species to elevated CO2. Aust. J. Plant Physiol., 27, 301-310.
Gremmen N.J.M., Chown S.L. & Marshall D.J. (1998). Impact of the introduced grass Agrostis
stolonifera on vegetation and soil fauna communities at Marion Island, sub-Antarctic. Biol.
Cons., 85, 223-231.
Grieco J.P., Johnson S., Achee N.L., Masuoka P., Pope K., Rejmankova E., Vanzie E., Andre R. &
Roberts D. (2006). Distribution of Anopheles albimanus, Anopheles vestitipennis, and
Anopheles crucians associated with land use in northern Belize. J. Med. Ent., 43, 614-622.
Griffen B.D. & Byers J.E. (2006). Intraguild predation reduces redundancy of predator species in
multiple predator assemblage. J. Anim. Ecol., 75, 959-966.
Grilli M.P. & Bruno M. (2007). Regional abundance of a planthopper pest: the effect of host patch area
and configuration. Ent. Exp. Appl., 122, 133-143.
Griswold M.W. & Lounibos L.P. (2005). Does differential predation permit invasive and native
mosquito larvae to coexist in Florida? Ecol. Ent., 30, 122-127.
Groenteman R., Guershon M. & Coll M. (2006). Effects of leaf nitrogen content on oviposition site
selection, offspring performance, and intraspecific interactions in an omnivorous bug. Ecol.
Ent., 31, 155-161.
Grogan P. & Chapin F.S. (2000). Nitrogen limitation of production in a Californian annual grassland:
The contribution of arbuscular mycorrhizae. Biogeochem., 49, 37-51.
Groom M.J. (2001). Consequences of subpopulation isolation for pollination, herbivory, and
population growth in Clarkia concinna concinna (Onagraceae). Biol. Cons., 100, 55-63.
Groppe K., Steinger T., Schmid B., Baur B. & Boller T. (2001). Effects of habitat fragmentation on
choke disease (Epichloe bromicola) in the grass Bromus erectus. J. Ecol., 89, 247-255.
Grosholz E.D. (2005). Recent biological invasion may hasten invasional meltdown by accelerating
historical introductions. Proc. Natl Acad. Sci. U.S.A., 102, 1088-1091.
Grosholz E.D., Ruiz G.M., Dean C.A., Shirley K.A., Maron J.L. & Connors P.G. (2000). The impacts
of a nonindigenous marine predator in a California bay. Ecology, 81, 1206-1224.
Gross C.L. (2001). The effect of introduced honeybees on native bee visitation and fruit-set in
Dillwynia juniperina (Fabaceae) in a fragmented ecosystem. Biol. Cons., 102, 89-95.
Gross C.L. & Mackay D. (1998). Honeybees reduce fitness in the pioneer shrub Melastoma affine
(Melastomataceae). Biol. Cons., 86, 169-178.
Grunzweig J.M. & Korner C. (2001). Biodiversity effects of elevated CO2 in species-rich model
communities from the semi-arid Negev of Israel. Oikos, 95, 112-124.
Guimarães P.R., Rico-Gray V., Oliveira P.S., Izzo T.J., dos Reis S.F. & Thompson J.N. (2007).
Interaction intimacy affects structure and coevolutionary dynamics in mutualistic networks.
Curr. Biol., 17, 1797-1803.
Haddad N.M., Haarstad J. & Tilman D. (2000). The effects of long-term nitrogen loading on grassland
insect communities. Oecologia, 124, 73-84.
Halpern S.L. & Underwood N. (2006). Approaches for testing herbivore effects on plant population
dynamics. J. Appl. Ecol., 43, 922-929.
Hamilton J.G., Dermody O., Aldea M., Zangerl A.R., Rogers A., Berenbaum M.R. & DeLucia E.H.
(2005). Anthropogenic changes in tropospheric composition increase susceptibility of soybean
to insect herbivory. Environ. Ent., 34, 479-485.
Hampton J.O., Spencer P.B.S., Alpers D.L., Twigg L.E., Woolnough A.P., Doust J., Higgs T. & Pluske
J. (2004). Molecular techniques, wildlife management and the importance of genetic
population structure and dispersal: a case study with feral pigs. J. Appl. Ecol., 41, 735-743.
Handa I.T., Korner C. & Hattenschwiler S. (2006). Conifer stem growth at the altitudinal treeline in
response to four years of CO 2 enrichment. Glob. Change Biol., 12, 2417-2430.
Hansen D.M., Olesen J.M. & Jones C.G. (2002). Trees, birds and bees in Mauritius: exploitative
competition between introduced honey bees and endemic nectarivorous birds? J. Biogeog., 29,
721-734.
Harrison R.D. (2000). Repercussions of El Niño: drought causes extinction and the breakdown of
mutualism in Borneo. Proc. R. Soc. Lond. B, 267, 911-915.
40
Harrison R.D. (2001). Drought and the consequences of El Niño in Borneo: a case study of figs. Pop.
Ecol., 43, 63-75.
Harte J., Rawa A. & Price V. (1996). Effects of manipulated soil microclimate on mesofaunal biomass
and diversity. Soil Biol. Biochem., 28, 313-322.
Hartley S.E., Gardner S.M. & Mitchell R.J. (2003). Indirect effects of grazing and nutrient addition on
the hemipteran community of heather moorlands. J. Appl. Ecol., 40, 793-803.
Hartley S.E. & Mitchell R.J. (2005). Manipulation of nutrients and grazing levels on heather moorland:
changes in Calluna dominance and consequences for community composition. J. Ecol., 93,
990-1004.
Hartwig U.A., Wittmann P., Raun R.B., Hartwig-Raz B., Jansa J., Mozafar A., Luscher A.,
Leuchtmann A., Frossard E. & Nosberger J. (2002). Arbuscular mycorrhiza infection
enhances the growth response of Lolium perenne to elevated atmospheric pCO2. J. Exp. Bot.,
53, 1207-1213.
Hattenschwiler S. & Schafellner C. (1999). Opposing effects of elevated CO2 and N deposition on
Lymantria monacha larvae feeding on spruce trees. Oecologia, 118, 210-217.
Hattenschwiler S. & Schafellner C. (2004). Gypsy moth feeding in the canopy of a CO2-enriched
mature forest. Glob. Change Biol., 10, 1899-1908.
Hattenschwiler S. & Zumbrunn T. (2006). Hemiparasite abundance in an alpine treeline ecotone
increases in response to atmospheric CO2 enrichment. Oecologia, 147, 47-52.
Hebeisen T., Luscher A., Zanetti S., Fischer B.U., Hartwig U.A., Frehner M., Hendrey G.R., Blum H.
& Nosberger J. (1997). Growth response of Trifolium repens L and Lolium perenne L as
monocultures and bi-species mixture to free air CO2 enrichment and management. Glob.
Change Biol., 3, 149-160.
Heegaard E. & Vandvik V. (2004). Climate change affects the outcome of competitive interactions - an
application of principal response curves. Oecologia, 139, 459-466.
Hegazi E. & Khafagi W. (2005). Developmental interaction between suboptimal instars of Spodoptera
littoralis (Lepidoptera : Noctuidae) and its parasitoid Microplitis rufiventris (Hymenoptera:
Braconidae). Arch. Insect Biochem. Physiol., 60, 172-184.
Heinemeyer A. & Fitter A.H. (2004). Impact of temperature on the arbuscular mycorrhizal (AM)
symbiosis: growth responses of the host plant and its AM fungal partner. J. Exp. Bot., 55, 525534.
Helgason T., Daniell T.J., Husband R., Fitter A.H. & Young J.P.W. (1998). Ploughing up the woodwide web? Nature, 394, 431-431.
Hellstrom K., Rautio P., Huhta A.P. & Tuomi J. (2004). Tolerance of an annual hemiparasite,
Euphrasia stricta agg., to simulated grazing in relation to the host environment. Flora, 199,
247-255.
Helms K.R. & Vinson S.B. (2005). Surface activity of native ants co-occurring with the red imported
fire ant, Solenopsis invicta (Hymenoptera : Formicidae). Southwest. Entomol., 30, 223-237.
Hely S.E.L. & Roxburgh S.H. (2005). The interactive effects of elevated CO2, temperature and initial
size on growth and competition between a native C-3 and an invasive C-3 grass. Plant Ecol.,
177, 85-98.
Hendrix P.F., Baker G.H., Callaham M.A., Damoff G.A., Fragoso C., Gonzalez G., James S.W.,
Lachnicht S.L., Winsome T. & Zou X. (2006). Invasion of exotic earthworms into ecosystems
inhabited by native earthworms. Biol. Inv., 8, 1287-1300.
Hendrix P.F., Parmelee R.W., Crossley D.A., Coleman D.C., Odum E.P. & Groffman P.M. (1986).
Detritus food webs in conventional and no-tillage agroecosystems. BioScience, 36, 374-380.
Hickerson C.A.M., Anthony C.D. & Walton B.M. (2005). Edge effects and intraguild predation in
native and introduced centipedes: evidence from the field and from laboratory microcosms.
Oecologia, 146, 110-119.
Hijri I., Sykorova Z., Oehl F., Ineichen K., Mader P., Wiemken A. & Redecker D. (2006).
Communities of arbuscular mycorrhizal fungi in arable soils are not necessarily low in
diversity. Molec. Ecol., 15, 2277-2289.
Hines J., Lynch M.E. & Denno R.F. (2005). Sap-feeding communities as indicators of habitat
fragmentation and nutrient subsidies. J. Insect Cons., 9, 261-280.
Hodkinson I.D. (2005). Terrestrial insects along elevation gradients: species and community responses
to altitude. Biol. Rev., 80, 489-513.
Hoeksema J.D., Lussenhop J. & Teeri J.A. (2000). Soil nematodes indicate food web responses to
elevated atmospheric CO2. Pedobiologia, 44, 725-735.
41
Holton M.K., Lindroth R.L. & Nordheim E.V. (2003). Foliar quality influences tree-herbivoreparasitoid interactions: effects of elevated CO2, O-3, and plant genotype. Oecologia, 137, 233244.
Honnay O., Jacquemyn H., Bossuyt B. & Hermy M. (2005). Forest fragmentation effects on patch
occupancy and population viability of herbaceous plant species. New Phytol., 166, 723-736.
Hooftman D.A.P., Billeter R.C., Schmid B. & Diemer M. (2004). Genetic effects of habitat
fragmentation on common species of Swiss fen meadows. Cons. Biol., 18, 1043-1051.
Hoogendoorn M. & Heimpel G.E. (2002). Indirect interactions between an introduced and a native
ladybird beetle species mediated by a shared parasitoid. Biol. Control, 25, 224-230.
Hoyle M. & Harborne A.R. (2005). Mixed effects of habitat fragmentation on species richness and
community structure in a microarthropod microecosystem. Ecol. Ent., 30, 684-691.
Hu S.J., Wu J.S., Burkey K.O. & Firestone M.K. (2005). Plant and microbial N acquisition under
elevated atmospheric CO2 in two mesocosm experiments with annual grasses. Glob. Change
Biol., 11, 213-223.
Hughes L. & Bazzaz F.A. (1997). Effect of elevated CO2 on interactions between the western flower
thrips, Frankliniella occidentalis (Thysanoptera: Thripidae) and the common milkweed,
Asclepias syriaca. Oecologia, 109, 286-290.
Hughes L. & Bazzaz F.A. (2001). Effects of elevated CO2 on five plant-aphid interactions. Ent. Exp.
Appl., 99, 87-96.
Hungate B.A., Jaeger C.H., Gamara G., Chapin F.S. & Field C.B. (2000). Soil microbiota in two
annual grasslands: responses to elevated atmospheric CO2. Oecologia, 124, 589-598.
Hutchinson T.C., Watmough S.A., Sager E.P.S. & Karagatzides J.D. (1998). Effects of excess nitrogen
deposition and soil acidification on sugar maple (Acer saccharum) in Ontario, Canada: an
experimental study. Can. J. For. Res., 28, 299-310.
Huxman T.E. & Smith S.D. (2001). Photosynthesis in an invasive grass and native forb at elevated CO 2
during an El Niño year in the Mojave Desert. Oecologia, 128, 193-201.
Hwangbo J.K., Seel W.E. & Woodin S.J. (2003). Short-term exposure to elevated atmospheric CO2
benefits the growth of a facultative annual root hemiparasite, Rhinanthus minor (L.), more
than that of its host, Poa pratensis (L.). J. Exp. Bot., 54, 1951-1955.
Ings T.C., Ward N.L. & Chittka L. (2006). Can commercially imported bumble bees out-compete their
native conspecifics? J. Appl. Ecol., 43, 940-948.
Ishtiaq F., Beadell J.S., Baker A.J., Rahmani A.R., Jhala Y.V. & Fleischer R.C. (2006). Prevalence and
evolutionary relationships of haematozoan parasites in native versus introduced populations of
common myna Acridotheres tristis. Proc. R. Soc. Lond. B, 273, 587-594.
Jacquemyn H., Brys R. & Hermy M. (2002). Patch occupancy, population size and reproductive
success of a forest herb (Primula elatior) in a fragmented landscape. Oecologia, 130, 617-625.
Jennersten O. (1988). Pollination in Dianthus deltoides (Caryophyllaceae): effects of habitat
fragmentation on visitation and seed set. Cons. Biol., 2, 359-66.
Jiang N. & Schulthess F. (2005). The effect of nitrogen fertilizer application to maize and sorghum on
the bionomics of Chilo partellus (Lepidoptera: Crambidae) and the performance of its larval
parasitoid Cotesia flavipes (Hymenoptera : Braconidae). Bull. Ent. Res., 95, 495-504.
Jifon J.L., Graham J.H., Drouillard D.L. & Syvertsen J.P. (2002). Growth depression of mycorrhizal
Citrus seedlings grown at high phosphorus supply is mitigated by elevated CO 2. New Phytol.,
153, 133-142.
Johns C.V., Beaumont L.J. & Hughes L. (2003). Effects of elevated CO2 and temperature on
development and consumption rates of Octotoma championi and O. scabripennis feeding on
Lantana camara. Ent. Exp. Appl., 108, 169-178.
Johns C.V. & Hughes A. (2002). Interactive effects of elevated CO2 and temperature on the leaf-miner
Dialectica scalariella Zeller (Lepidoptera : Gracillariidae) in Paterson's Curse, Echium
plantagineum (Boraginaceae). Glob. Change Biol., 8, 142-152.
Johnson D., Leake J.R., Lee J.A. & Campbell C.D. (1998). Changes in soil microbial biomass and
microbial activities in response to 7 years simulated pollutant nitrogen deposition on a
heathland and two grasslands. Environ. Pollution, 103, 239-250.
Johnson D., Leake J.R. & Read D.J. (2005a). Liming and nitrogen fertilization affects phosphatase
activities, microbial biomass and mycorrhizal colonisation in upland grassland. Plant Soil,
271, 157-164.
Johnson M.T., Follett P.A., Taylor A.D. & Jones V.P. (2005b). Impacts of biological control and
invasive species on a non-target native Hawaiian insect. Oecologia, 142, 529-540.
42
Johnson N.C., Rowland D.L., Corkidi L., Egerton-Warburton L.M. & Allen E.B. (2003). Nitrogen
enrichment alters mycorrhizal allocation at five mesic to semiarid grasslands. Ecology, 84,
1895-1908.
Johnson N.C., Wolf J., Reyes M.A., Panter A., Koch G.W. & Redman A. (2005c). Species of plants
and associated arbuscular mycorrhizal fungi mediate mycorrhizal responses to CO 2
enrichment. Glob. Change Biol., 11, 1156-1166.
Johnson S.D., Collin C.L., Wissman H.J., Halvarsson E. & Agren J. (2004a). Factors contributing to
variation in seed production among remnant populations of the endangered daisy Gerbera
aurantiaca. Biotropica, 36, 148-155.
Johnson S.D., Neal P.R., Peter C.I. & Edwards T.J. (2004b). Fruiting failure and limited recruitment in
remnant populations of the hawkmoth-pollinated tree Oxyanthus pyriformis subsp. pyriformis
(Rubiaceae). Biol. Cons., 120, 31-39.
Jones T.H., Thompson L.J., Lawton J.H., Bezemer T.M., Bardgett R.D., Blackburn T.M., Bruce K.D.,
Cannon P.F., Hall G.S., Hartley S.E., Howson G., Jones C.G., Kampichler C., Kandeler E. &
Ritchie D.A. (1998). Impacts of rising atmospheric carbon dioxide on model terrestrial
ecosystems. Science, 280, 441-443.
Jordano P. (2000). Fruits and frugivory. In: Seeds: the Ecology of Regeneration in Plant Communities.
2nd edition (ed. Fenner M). CABI Wallingford, U.K., pp. 125-166.
Ju H.J., Hill N.S., Abbott I. & Ingram K.T. (2006). Temperature influences on endophyte growth in tall
fescue. Crop Sci., 46, 404-412.
Jules E.S., Kauffman M.J., Ritts W.D. & Carroll A.L. (2002). Spread of an invasive pathogen over a
variable landscape: A nonnative root rot on Port Orford cedar. Ecology, 83, 3167-3181.
Jules E.S. & Rathcke B.J. (1999). Mechanisms of reduced Trillium recruitment along edges of oldgrowth forest fragments. Cons. Biol., 13, 784-793.
Jwa N.S. & Walling L.L. (2001). Influence of elevated CO2 concentration on disease development in
tomato. New Phytol., 149, 509-518.
Kamata N., Esaki K., Kato K., Igeta Y. & Wada K. (2002). Potential impact of global warming on
deciduous oak dieback caused by ambrosia fungus Raffaelea sp carried by ambrosia beetle
Platypus quercivorus (Coleoptera : Platypodidae) in Japan. Bull.Ent.Res., 92, 119-126.
Kandeler E., Tscherko D., Bardgett R.D., Hobbs P.J., Kampichler C. & Jones T.H. (1998). The
response of soil microorganisms and roots to elevated CO2 and temperature in a terrestrial
model ecosystem. Plant Soil, 202, 251-262.
Kardol P., Bezemer T.M., van der Wal A. & van der Putten W.H. (2005). Successional trajectories of
soil nematode and plant communities in a chronosequence of ex-arable lands. Biol. Cons.,
126, 317-327.
Karrenberg S. & Jensen K. (2000). Effects of pollination and pollen source on the seed set of
Pedicularis palustris. Folia Geobot., 35, 191-202.
Kaspari M. (2005). Global energy gradients and size in colonial organisms: Worker mass and worker
number in ant colonies. Proc. Natl Acad. Sci. U.S.A., 102, 5079-5083.
Kasurinen A., Helmisaari H.S. & Holopainen T. (1999). The influence of elevated CO2 and O-3 on fine
roots and mycorrhizas of naturally growing young Scots pine trees during three exposure
years. Glob. Change Biol., 5, 771-780.
Kato M., Shibata A., Yasui T. & Nagamasu H. (1999). Impact of introduced honeybees, Apis mellifera,
upon native bee communities in the Bonin (Ogasawara) Islands. Res. Pop. Ecol., 41, 217-228.
Kaye J.P. & Hart S.C. (1997). Competition for nitrogen between plants and soil microorganisms.
Trends Ecol. Evol., 12, 139-143.
Keeler M.S., Chew F.S., Goodale B.C. & Reed J.M. (2006). Modelling the impacts of two exotic
invasive species on a native butterfly: top-down vs. bottom-up effects. J. Anim. Ecol., 75, 777788.
Kelly D., Ladley J.J., Robertson A.W. & Norton D.A. (2000). Limited forest fragmentation improves
reproduction in the declining New Zealand mistletoe Peraxilla tetrapetala (Loranthaceae). In:
Genetics, Demography and Viability of Fragmented Populations (eds. Young AG & Clarke
G). Cambridge University Press Cambridge, UK, pp. 241-252.
Kelly D., Robertson A.W., Ladley J.J., Anderson S.H. & McKenzie R.J. (2006). Relative
(un)importance of introduced animals as pollinators and dispersers of native plants. In:
Biological Invasions in New Zealand (eds. Allen RB & Lee WG). Springer Berlin.
Kelt D.A., Meserve P.L., Nabors L.K., Forister M.L. & Gutierrez J.R. (2004). Foraging ecology of
small mammals in semiarid Chile: The interplay of biotic and abiotic effects. Ecology, 85,
383-397.
43
Kerslake J.E., Woodin S.J. & Hartley S.E. (1998). Effects of carbon dioxide and nitrogen enrichment
on a plant-insect interaction: the quality of Calluna vulgaris as a host for Operophtera
brumata. New Phytol., 140, 43-53.
Kiesecker J.M., Blaustein A.R. & Miller C.L. (2001). Transfer of a pathogen from fish to amphibians.
Cons. Biol., 15, 1064-1070.
King J.R. & Tschinkel W.R. (2006). Experimental evidence that the introduced fire ant, Solenopsis
invicta, does not competitively suppress co-occurring ants in a disturbed habitat. J. Anim.
Ecol., 75, 1370-1378.
Kiritani K. (2007). The impact of global warming and land-use change on the pest status of rice and
fruit bugs (Heteroptera) in Japan. Glob. Change Biol., 13, 1586-1595.
Klanderud K. (2005). Climate change effects on species interactions in an alpine plant community. J.
Ecol., 93, 127-137.
Klein A.-M., Steffan-Dewenter I. & Tscharntke T. (2003a). Fruit set of highland coffee increases with
the diversity of pollinating bees. Proc. R. Soc. Lond. Ser. B-Biol. Sci., 270, 955-961.
Klein A.M., Steffan-Dewenter I. & Tscharntke T. (2002). Predator-prey ratios on cocoa along a landuse gradient in Indonesia. Biodiv. Conserv., 11, 683-693.
Klein A.M., Steffan-Dewenter I. & Tscharntke T. (2003b). Pollination of Coffea canephora in relation
to local and regional agroforestry management. J. Appl. Ecol., 40, 837-845.
Klein A.M., Vaissiere B.E., Cane J.H., Steffan-Dewenter I., Cunningham S.A., Kremen C. &
Tscharntke T. (2007). Importance of pollinators in changing landscapes for world crops. Proc.
R. Soc. Lond. B, 274, 303-313.
Klein D.A., Paschke M.W. & Heskett T.L. (2006). Comparative fungal responses in managed plant
communities infested by spotted (Centaurea maculosa Lam.) and diffuse (C. diffusa Lam.)
knapweed. Appl. Soil Ecol., 32, 89-97.
Klironomos J.N., Rillig M.C. & Allen M.F. (1996). Below-ground microbial and microfaunal
responses to Artemisia tridentata grown under elevated atmospheric CO2. Funct. Ecol., 10,
527-534.
Klironomos J.N., Rillig M.C., Allen M.F., Zak D.R., Kubiske M. & Pregitzer K.S. (1997a). Soil
fungal-arthropod responses to Populus tremuloides grown under enriched atmospheric CO 2
under field conditions. Glob. Change Biol., 3, 473-478.
Klironomos J.N., Rillig M.C., Allen M.F., Zak D.R., Pregitzer K.S. & Kubiske M.E. (1997b).
Increased levels of airborne fungal spores in response to Populus tremuloides grown under
elevated atmospheric CO2. Can. J. Bot.-Rev. Can. Bot., 75, 1670-1673.
Klironomos J.N., Ursic M., Rillig M. & Allen M.F. (1998). Interspecific differences in the response of
arbuscular mycorrhizal fungi to Artemisia tridentata grown under elevated atmospheric CO2.
New Phytol., 138, 599-605.
Knepp R.G., Hamilton J.G., Mohan J.E., Zangerl A.R., Berenbaum M.R. & DeLucia E.H. (2005).
Elevated CO2 reduces leaf damage by insect herbivores in a forest community. New Phytol.,
167, 207-218.
Knox M.R., Torres-Acosta J.F.J. & Aguilar-Caballero A.J. (2006). Exploiting the effect of dietary
supplementation of small ruminants on resilience and resistance against gastrointestinal
nematodes. Vet. Parasitol., 139, 385-393.
Koenig W.D. (2003). European starlings and their effect on native cavity-nesting birds. Cons. Biol., 17,
1134-1140.
Kolb A. (2005). Reduced reproductive success and offspring survival in fragmented populations of the
forest herb Phyteuma spicatum. J. Ecol., 93, 1226-1237.
Kollmann J., Coomes D.A. & White S.M. (1998). Consistencies in post-dispersal seed predation of
temperate fleshy-fruited species among seasons, years and sites. Funct. Ecol., 12, 683-690.
Kourtev P.S., Ehrenfeld J.G. & Haggblom M. (2003). Experimental analysis of the effect of exotic and
native plant species on the structure and function of soil microbial communities. Soil Biol.
Biochem., 35, 895-905.
Krauss J., Harri S.A., Bush L., Husi R., Bigler L., Power S.A. & Muller C.B. (2007). Effects of
fertilizer, fungal endophytes and plant cultivar on the performance of insect herbivores and
their natural enemies. Funct. Ecol., 21, 107-116.
Kremen C., Williams N.M. & Thorp R.W. (2002). Crop pollination from native bees at risk from
agricultural intensification. Proc. Natl Acad. Sci. U.S.A., 99, 16812-16816.
Kruess A. (2003). Effects of landscape structure and habitat type on a plant-herbivore-parasitoid
community. Ecography, 26, 283-290.
Kruess A. & Tscharntke T. (1994). Habitat fragmentation, species loss, and biological control. Science,
264, 1581-1584.
44
Kruess A. & Tscharntke T. (2000). Species richness and parasitism in a fragmented landscape:
Experiments and field studies with insects on Vicia sepium. Oecologia, 122, 129-137.
Kudo G. & Hirao A.S. (2006). Habitat-specific responses in the flowering phenology and seed set of
alpine plants to climate variation: implications for global-change impacts. Pop. Ecol., 48, 4958.
Kuijper D.P.J., Dubbeld J. & Bakker J.P. (2005). Competition between two grass species with and
without grazing over a productivity gradient. Plant Ecol., 179, 237-246.
Kutz S.J., Hoberg E.P., Polley L. & Jenkins E.J. (2005). Global warming is changing the dynamics of
Arctic host-parasite systems. Proc. R. Soc. Lond. B, 272, 2571-2576.
Lacey E.P., Roach D.A., Herr D., Kincaid S. & Perrott R. (2003). Multigenerational effects of
flowering and fruiting phenology in Plantago lanceolata. Ecology, 84, 2462-2475.
Lake J.C. & Hughes L. (1999). Nectar production and floral characteristics of Tropaeolum majus L.
grown in ambient and elevated carbon dioxide. Ann. Bot., 84, 535-541.
Lambrecht S.C., Loik M.E., Inouye D.W. & Harte J. (2007). Reproductive and physiological responses
to simulated climate warming for four subalpine species. New Phytol., 173, 121-134.
Lamont B.B., Klinkhamer P.G.L. & Witkowski E.T.F. (1993). Population fragmentation may reduce
fertility to zero in Banksia goodii - a demonstration of the Allee effect. Oecologia, 94, 446450.
Lange O.L. & Green T.G.A. (2005). Lichens show that fungi can acclimate their respiration to seasonal
changes in temperature. Oecologia, 142, 11-19.
Langley J.A., Dijkstra P., Drake B.G. & Hungate B.A. (2003). Ectomycorrhizal colonization, biomass,
and production in a regenerating scrub oak forest in response to elevated CO 2. Ecosyst., 6,
424-430.
Larson D.L., Royer R.A. & Royer M.R. (2006). Insect visitation and pollen deposition in an invaded
prairie plant community. Biol. Cons., 130, 148-159.
Lau J.A. & Strauss S.Y. (2005). Insect herbivores drive important indirect effects of exotic plants on
native communities. Ecology, 86, 2990-2997.
Laurance W.F., Nascimento H.E.M., Laurance S.G., Andrade A., Ribeiro J., Giraldo J.P., Lovejoy
T.E., Condit R., Chave J., Harms K.E. & D'Angelo S. (2006). Rapid decay of tree-community
composition in Amazonian forest fragments. Proc. Natl Acad. Sci. U.S.A., 103, 19010-19014.
Laurance W.F., Oliveira A.A., Laurance S.G., Condit R., Nascimento H.E.M., Sanchez-Thorin A.C.,
Lovejoy T.E., Andrade A., D'Angelo S., Ribeiro J.E. & Dick C.W. (2004). Pervasive
alteration of tree communities in undisturbed Amazonian forests. Nature, 428, 171-175.
Laurance W.F. & Williamson G.B. (2001). Positive feedbacks among forest fragmentation, drought,
and climate change in the Amazon. Cons. Biol., 15, 1529-1535.
Lavorel S., McIntyre S., Landsberg J. & Forbes T.D.A. (1997). Plant functional classifications: from
general groups to specific groups based on response to disturbance. Trends Ecol. Evol., 12,
474-478.
Lazaro A., Traveset A. & Mendez M. (2006). Masting in Buxus balearica: assessing fruiting patterns
and processes at a large spatial scale. Oikos, 115, 229-240.
Leimu R. & Syrjanen K. (2002). Effects of population size, seed predation and plant size on male and
female reproductive success in Vincetoxicum hirundinaria (Asclepiadaceae). Oikos, 98, 229238.
Lejeune K.D., Suding K.N., Sturgis S., Scott A. & Seastedt T.R. (2005). Biological control insect use
of fertilized and unfertilized diffuse knapweed in a Colorado grassland. Environ. Ent., 34,
225-234.
Lennartsson T. (2002). Extinction thresholds and disrupted plant-pollinator interactions in fragmented
plant populations. Ecology, 83, 3060-3072.
Lenz L. & Taylor J.A. (2001). The influence of an invasive tree species (Myrica faya) on the
abundance of an alien insect (Sophonia rufofascia) in Hawai'i Volcanoes National Park. Biol.
Cons., 102, 301-307.
Levin P.S., Coyer J.A., Petrik R. & Good T.P. (2002). Community-wide effects of nonindigenous
species on temperate rocky reefs. Ecology, 83, 3182-3193.
Lewinsohn T.M., Prado P.I., Jordano P., Bascompte J. & Olesen J.M. (2006). Structure in plant-animal
interaction assemblages. Oikos, 113, 174-184.
Lewis J.D., Thomas R.B. & Strain B.R. (1994). Effect of elevated CO2 on mycorrhizal colonization of
Lolloby-pine (Pinus taeda L) seedlings. Plant Soil, 165, 81-88.
Li Q., Liang W.J., Jiang Y., Shi Y., Zhu J.G. & Neher D.A. (2007). Effect of elevated CO2 and N
fertilisation on soil nematode abundance and diversity in a wheat field. Appl. Soil Ecol., 36,
63-69.
45
Li W.H., Zhang C.B., Jiang H.B., Xin G.R. & Yang Z.Y. (2006). Changes in soil microbial community
associated with invasion of the exotic weed, Mikania micrantha HBK. Plant Soil, 281, 309324.
Lienert J. & Fischer M. (2003). Habitat fragmentation affects the common wetland specialist Primula
farinosa in north-east Switzerland. J. Ecol., 91, 587-599.
Lilleskov E.A., Fahey T.J., Horton T.R. & Lovett G.M. (2002). Belowground ectomycorrhizal fungal
community change over a nitrogen deposition gradient in Alaska. Ecology, 83, 104-115.
Lilleskov E.A., Fahey T.J. & Lovett G.M. (2001). Ectomycorrhizal fungal aboveground community
change over an atmospheric nitrogen deposition gradient. Ecol. Appl., 11, 397-410.
Lilley J.H., Cerenius L. & Soderhall K. (1997). RAPD evidence for the origin of crayfish plague
outbreaks in Britain. Aquaculture, 157, 181-185.
Limpens J., Raymakers J., Baar J., Berendse F. & Zijlstra J.D. (2003). The interaction between
epiphytic algae, a parasitic fungus and Sphagnum as affected by N and P. Oikos, 103, 59-68.
Lindgren E. & Gustafson R. (2001). Tick-borne encephalitis in Sweden and climate change. Lancet,
358, 16-18.
Liu H. & Stiling P. (2006). Testing the enemy release hypothesis: a review and meta-analysis. Biol.
Inv., 8, 1535-1545.
Logan J.A., Regniere J. & Powell J.A. (2003). Assessing the impacts of global warming on forest pest
dynamics. Front. Ecol. Environ., 1, 130-137.
LoGiudice K., Ostfeld R.S., Schmidt K.A. & Keesing F. (2003). The ecology of infectious disease:
Effects of host diversity and community composition on Lyme disease risk. Proc. Natl Acad.
Sci. U.S.A., 100, 567-571.
Long S.P., Ainsworth E.A., Leakey A.D.B., Nosberger J. & Ort D.R. (2006). Food for thought: Lowerthan-expected crop yield stimulation with rising CO2 concentrations. Science, 312, 1918-1921.
Lou Y.G. & Baldwin I.T. (2004). Nitrogen supply influences herbivore-induced direct and indirect
defenses and transcriptional responses to Nicotiana attenuata. Plant Physiology, 135, 496506.
Louche-Tessandier D., Samson G., Hernandez-Sebastia C., Chagvardieff P. & Desjardins Y. (1999).
Importance of light and CO2 on the effects of endomycorrhizal colonization on growth and
photosynthesis of potato plantlets (Solanum tuberosum) in an in vitro tripartite system. New
Phytol., 142, 539-550.
Lovelock C.E., Winter K., Mersits R. & Popp M. (1998). Responses of communities of tropical tree
species to elevated CO2 in a forest clearing. Oecologia, 116, 207-218.
Luijten S.H., Dierick A., Oostermeijer J.G.B., Raijmann L.E.L. & Den Nijs H.C.M. (2000). Population
size, genetic variation, and reproductive success in a rapidly declining, self-incompatible
perennial (Arnica montana) in The Netherlands. Cons. Biol., 14, 1776-1787.
Lukac M., Calfapietra C. & Godbold D.L. (2003). Production, turnover and mycorrhizal colonization
of root systems of three Populus species grown under elevated CO2 (POPFACE). Glob.
Change Biol., 9, 838-848.
Lussenhop J., Treonis A., Curtis P.S., Teeri J.A. & Vogel C.S. (1998). Response of soil biota to
elevated atmospheric CO2 in poplar model systems. Oecologia, 113, 247-251.
Maerz J.C., Karuzas J.M., Madison D.M. & Blossey B. (2005). Introduced invertebrates are important
prey for a generalist predator. Divers. Distrib., 11, 83-90.
Mahoro S. (2003). Effects of flower and seed predators and pollinators on fruit production in two
sequentially flowering congeners. Plant Ecol., 166, 37-48.
Malmstrom C.M., McCullough A.J., Johnson H.A., Newton L.A. & Borer E.T. (2005). Invasive annual
grasses indirectly increase virus incidence in California native perennial bunchgrasses.
Oecologia, 145, 153-164.
Malmstrom C.M., Stoner C.J., Brandenburg S. & Newton L.A. (2006). Virus infection and grazing
exert counteracting influences on survivorship of native bunchgrass seedlings competing with
invasive exotics. J. Ecol., 94, 264-275.
Mangan S.A., Eom A.H., Adler G.H., Yavitt J.B. & Herre E.A. (2004). Diversity of arbuscular
mycorrhizal fungi across a fragmented forest in Panama: insular spore communities differ
from mainland communities. Oecologia, 141, 687-700.
Manning P., Newington J.E., Robson H.R., Saunders M., Eggers T., Bradford M.A., Bardgett R.D.,
Bonkowski M., Ellis R.J., Gange A.C., Grayston S.J., Kandeler E., Marhan S., Reid E.,
Tscherko D., Godfray H.C.J. & Rees M. (2006). Decoupling the direct and indirect effects of
nitrogen deposition on ecosystem function. Ecol. Lett., 9, 1015-1024.
Marks S. & Lincoln D.E. (1996). Antiherbivore defense mutualism under elevated carbon dioxide
levels: A fungal endophyte and grass. Environ. Ent., 25, 618-623.
46
Marschner P., Grierson P.F. & Rengel Z. (2005). Microbial community composition and functioning in
the rhizosphere of three Banksia species in native woodland in Western Australia. Appl. Soil
Ecol., 28, 191-201.
Martin T.E. (2007). Climate correlates of 20 years of trophic changes in a high-elevation riparian
system. Ecology, 88, 367-380.
Mathimaran N., Ruh R., Jama B., Verchot L., Frossard E. & Jansa J. (2007). Impact of agricultural
management on arbuscular mycorrhizal fungal communities in Kenyan ferralsol. Agric.
Ecosyst. Environ., 119, 22-32.
Matthies D. & Egli P. (1999). Response of a root hemiparasite to elevated CO2 depends on host type
and soil nutrients. Oecologia, 120, 156-161.
Mavraganis K. & Eckert C.G. (2001). Effects of population size and isolation on reproductive output in
Aquilegia canadensis (Ranunculaceae). Oikos, 95, 300-310.
McDonald R.A., O'Hara K. & Morrish D.J. (2007). Decline of invasive alien mink (Mustela vison) is
concurrent with recovery of native otters (Lutra lutra). Divers. Distrib., 13, 92-98.
McEuen A.B. & Curran L.M. (2006). Plant recruitment bottlenecks in temperate forest fragments: seed
limitation and insect herbivory. Plant Ecol., 184, 297-309.
McKone M.J., Kelly D. & Lee W.G. (1998). Effect of climate change on mast-seeding species:
frequency of mass flowering and escape from specialist insect seed predators. Glob. Change
Biol., 4, 591-596.
McLean J.A. & Leckie A.C. (2000). Vespula vulgaris (Hymenoptera: Vespidae) dominates December
malaise trap catches in a West Coast honeydew beech forest. N.Z. Ent., 22, 69-72.
McLean M. & Parkinson D. (1998a). Impacts of the epigeic earthworm Dendrobaena octaedra on
oribatid mite community diversity and microarthropod abundances in pine forest floor: a
mesocosm study. Appl. Soil Ecol., 7, 125-136.
McLean M.A., Migge-Kleian S. & Parkinson D. (2006). Earthworm invasions of ecosystems devoid of
earthworms: effects on soil microbes. Biol. Inv., 8, 1257-1273.
McLean M.A. & Parkinson D. (1998b). Impacts of the epigeic earthworm Dendrobaena octaedra on
microfungal community structure in pine forest floor: a mesocosm study. Appl. Soil Ecol., 8,
61-75.
McLean M.A. & Parkinson D. (2000). Field evidence of the effects of the epigeic earthworm
Dendrobaena octaedra on the microfungal community in pine forest floor. Soil Biol.
Biochem., 32, 351-360.
Meijer G. & Leuchtmann A. (2000). The effects of genetic and environmental factors on disease
expression (stroma formation) and plant growth in Brachypodium sylvaticum infected by
Epichloe sylvatica. Oikos, 91, 446-458.
Memmott J., Craze P., Waser N. & Price M. (2007). Global warming and the disruption of plantpollinator interactions. Ecol. Lett., 10, 710-717.
Memmott J. & Waser N.M. (2002). Integration of alien plants into a native flower-pollinator visitation
web. Proc. R. Soc. Lond. B, 269, 2395-2399.
Mevi-Schutz J. & Erhardt A. (2005). Amino acids in nectar enhance butterfly fecundity: A longawaited link. Am. Nat., 165, 411-419.
Mevi-Schutz J., Goverde M. & Erhardt A. (2003). Effects of fertilization and elevated CO 2 on larval
food and butterfly nectar amino acid preference in Coenonympha pamphilus L. Behav. Ecol.
Sociobiol., 54, 36-43.
Mills L.S. (1995). Edge effects and isolation: Red-Backed Voles on forest remnants. Cons. Biol., 9,
395-403.
Mitchell C.E., Agrawal A.A., Bever J.D., Gilbert G.S., Hufbauer R.A., Klironomos J.N., Maron J.L.,
Morris W.F., Parker I.M., Power A.G., Seabloom E.W., Torchin M.E. & Vazquez D.P.
(2006). Biotic interactions and plant invasions. Ecol. Lett., 9, 726-740.
Mitchell C.E., Reich P.B., Tilman D. & Groth J.V. (2003). Effects of elevated CO2, nitrogen
deposition, and decreased species diversity on foliar fungal plant disease. Glob. Change Biol.,
9, 438-451.
Montgomery B.R., Kelly D., Robertson A.W. & Ladley J.J. (2003). Pollinator behaviour, not increased
resources, boosts seed set on forest edges in a New Zealand Loranthaceous mistletoe. New
Zealand Journal of Botany, 41, 277-286.
Moody-Weis J.M. & Heywood J.S. (2001). Pollination limitation to reproductive success in the
Missouri evening primrose, Oenothera macrocarpa (Onagraceae). Am. J. Bot., 88, 1615-1622.
Moon D.C., Rossi A.M. & Stiling P. (2000). The effects of abiotically induced changes in host plant
quality (and morphology) on a salt marsh planthopper and its parasitoid. Ecol. Ent., 25, 325331.
47
Moon D.C. & Stiling P. (2000). Relative importance of abiotically induced direct and indirect effects
on a salt-marsh herbivore. Ecology, 81, 470-481.
Moragues E. & Traveset A. (2005). Effect of Carpobrotus spp. on the pollination success of native
plant species of the Balearic Islands. Biol. Cons., 122, 611-619.
Moran M.D., Rooney T.P. & Hurd L.E. (1996). Top-down cascade from a bitrophic predator in an oldfield community. Ecology, 77, 2219-27.
Morgan J.W. (1999). Effects of population size on seed production and germinability in an endangered,
fragmented grassland plant. Cons. Biol., 13, 266-273.
Morrison L.W. (2002). Long-term impacts of an arthropod-community invasion by the imported fire
ant, Solenopsis invicta. Ecology, 83, 2337-2345.
Mouritsen K.N., Tompkins D.M. & Poulin R. (2005). Climate warming may cause a parasite-induced
collapse in coastal amphipod populations. Oecologia, 146, 476-483.
Mulder C., De Zwart D., Van Wijnen H.J., Schouten A.J. & Breure A.M. (2003). Observational and
simulated evidence of ecological shifts within the soil nematode community of
agroecosystems under conventional and organic farming. Funct. Ecol., 17, 516-525.
Muller C.B., Fellowes M.D.E. & Godfray H.C.J. (2005). Relative importance of fertiliser addition to
plants and exclusion of predators for aphid growth in the field. Oecologia, 143, 419-427.
Mummey D.L. & Rillig M.C. (2006). The invasive plant species Centaurea maculosa alters arbuscular
mycorrhizal fungal communities in the field. Plant Soil, 288, 81-90.
Muñoz A., Celedon-Neghme C., Cavieres L.A. & Arroyo M.T.K. (2005). Bottom-up effects of nutrient
availability on flower production, pollinator visitation, and seed output in a high-Andean
shrub. Oecologia, 143, 126-135.
Musolin D.L. (2007). Insects in a warmer world: ecological, physiological and life-history responses of
true bugs (Heteroptera) to climate change. Glob. Change Biol., 13, 1565-1585.
Mustajarvi K., Siikamaki P., Rytkonen S. & Lammi A. (2001). Consequences of plant population size
and density for plant-pollinator interactions and plant performance. J. Ecol., 89, 80-87.
Navas M.L., Garnier E., Austin M.P. & Gifford R.M. (1999). Effect of competition on the responses of
grasses and legumes to elevated atmospheric CO2 along a nitrogen gradient: differences
between isolated plants, monocultures and multi-species mixtures. New Phytol., 143, 323-331.
Neher D.A., Weicht T.R., Moorhead D.L. & Sinsabaugh R.L. (2004). Elevated CO2 alters functional
attributes of nematode communities in forest soils. Funct. Ecol., 18, 584-591.
Ness J.H. & Bressmer K. (2005). Abiotic influences on the behaviour of rodents, ants, and plants affect
an ant-seed mutualism. EcoSci., 12, 76-81.
Ness J.H., Bronstein J.L., Andersen A.N. & Holland J.N. (2004). Ant body size predicts dispersal
distance of ant-adapted seeds: Implications of small-ant invasions. Ecology, 85, 1244-1250.
Neuhauser C. & Fargione J.E. (2004). A mutualisim-parasitism continuum model and its application to
plant-mycorrhizae interactions. Ecol. Model., 177, 337-352.
Nielsen A. & Ims R.A. (2000). Bumble bee pollination of the sticky catchfly in a fragmented
agricultural landscape. EcoSci., 7, 157-165.
Niemala J., Spence J.R. & Carcamo H. (1997). Establishment and interactions of carabid populations:
an experiment with native and introduced species. Ecography, 20, 643-652.
Niklaus P.A., Alphei D., Ebersberger D., Kampichler C., Kandeler E. & Tscherko D. (2003). Six years
of in situ CO2 enrichment evoke changes in soil structure and soil biota of nutrient-poor
grassland. Glob. Change Biol., 9, 585-600.
Niklaus P.A., Wohlfender M., Siegwolf R. & Korner C. (2001). Effects of six years atmospheric CO2
enrichment on plant, soil, and soil microbial C of a calcareous grassland. Plant Soil, 233, 189202.
Noonburg E.G. & Byers J.E. (2005). More harm than good: When invader vulnerability to predators
enhances impact on native species. Ecology, 86, 2555-2560.
Norbury G. (2001). Conserving dryland lizards by reducing predator-mediated apparent competition
and direct competition with introduced rabbits. J. Appl. Ecol., 38, 1350-1361.
Nordin A., Nasholm T. & Ericson L. (1998). Effects of simulated N deposition on understorey
vegetation of a boreal coniferous forest. Funct. Ecol., 12, 691-699.
O'Dowd D.J., Green P.T. & Lake P.S. (2003). Invasional 'meltdown' on an oceanic island. Ecol. Lett.,
6, 812-817.
Ockinger E. & Smith H.G. (2007). Semi-natural grasslands as population sources for pollinating insects
in agricultural landscapes. J. Appl. Ecol., 44, 50-59.
Ogden N.H., Maarouf A., Barker I.K., Bigras-Poulin M., Lindsay L.R., Morshed M.G., O'Callaghan
C.J., Ramay F., Waltner-Toews D. & Charron D.F. (2006). Climate change and the potential
48
for range expansion of the Lyme disease vector Ixodes scapularis in Canada. Intl J. Parasitol.,
36, 63-70.
Olesen J.M., Bascompte J., Elberling H. & Jordano P. (2008). Temporal dynamics in a pollination
network. Ecology, 89, 1573-1582.
Olesniewicz K.S. & Thomas R.B. (1999). Effects of mycorrhizal colonization on biomass production
and nitrogen fixation of black locust (Robinia pseudoacacia) seedlings grown under elevated
atmospheric carbon dioxide. New Phytol., 142, 133-140.
Oostermeijer J.G.B. & van Swaay C.A.M. (1998). The relationship between butterflies and
environmental indicator values: a tool for conservation in a changing landscape. Biol. Cons.,
86, 271-280.
Opik M., Moora M., Liira J. & Zobel M. (2006). Composition of root-colonizing arbuscular
mycorrhizal fungal communities in different ecosystems around the globe. J. Ecol., 94, 778790.
Orrock J.L. & Damschen E.I. (2005). Corridors cause differential seed predation. Ecol. Appl., 15, 793798.
Ortega Y.K., Pearson D.E. & McKelvey K.S. (2004). Effects of biological control agents and exotic
plant invasion on deer mouse populations. Ecol. Appl., 14, 241-253.
Osborne J.L., Awmack C.S., Clark S.J., Williams I.H. & Mills V.C. (1997). Nectar and flower
production in Vicia faba L (field bean) at ambient and elevated carbon dioxide. Apidologie,
28, 43-55.
Ostergard H. & Ehrlen J. (2005). Among population variation in specialist and generalist seed
predation - the importance of host plant distribution, alternative hosts and environmental
variation. Oikos, 111, 39-46.
Pacini E., Nepi M. & Vesprini J.L. (2003). Nectar biodiversity: a short review. Plant Syst. Evol., 238,
7-21.
Pakkala T., Kouki J. & Tainen J. (2006). Top predator and interference competition modify the
occurrence and breeding success of a specialist species in a structurally complex forest
environment. Ann. Zool. Fennici, 43, 137-164.
Parmesan C. (2006). Ecological and evolutionary responses to recent climate change. Ann. Rev. Ecol.
Evol. Syst., 37, 637-669.
Parra-Tabla V., Vargas C.F., Magaña-Rueda S. & Navarro J. (2000). Female and male pollination
success of Oncidium ascendens Lindey (Orchidaceae) in two contrasting habitat patches:
forest vs agricultural field. Biol. Cons., 94, 335-340.
Parrent J.L., Morris W.F. & Vilgalys R. (2006). CO2-enrichment and nutrient availability alter
ectomycorrhizal fungal communities. Ecology, 87, 2278-2287.
Paschke M., Abs C. & Schmid B. (2002). Effects of population size and pollen diversity on
reproductive success and offspring size in the narrow endemic Cochlearia bavarica
(Brassicaceae). Am. J. Bot., 89, 1250-1259.
Patterson D.T., Westbrook J.K., Joyce R.J.V., Lingren P.D. & Rogasik J. (1999). Weeds, insects, and
diseases. Climatic Change, 43, 711-727.
Pearson D.E. & Callaway R.M. (2003). Indirect effects of host-specific biological control agents.
Trends Ecol. Evol., 18, 456-461.
Pearson D.E., McKelvey K.S. & Ruggiero L.F. (2000). Non-target effects of an introduced biological
control agent on deer mouse ecology. Oecologia, 122, 121-128.
Peay K.G., Bruns T.D., Kennedy P.G., Bergemann S.E. & Garbelotto M. (2007). A strong species-area
relationship for eukaryotic soil microbes: island size matters for ectomycorrhizal fungi. Ecol.
Lett., 10, 470-480.
Percy K.E., Awmack C.S., Lindroth R.L., Kubiske M.E., Kopper B.J., Isebrands J.G., Pregitzer K.S.,
Hendrey G.R., Dickson R.E., Zak D.R., Oksanen E., Sober J., Harrington R. & Karnosky D.F.
(2002). Altered performance of forest pests under atmospheres enriched by CO 2 and O-3.
Nature, 420, 403-407.
Perdikis D.C., Lykouressis D.P. & Economou L.P. (1999). The influence of temperature, photoperiod
and plant type on the predation rate of Macrolophus pygmaeus on Myzus persicae. Biocontrol,
44, 281-289.
Petanidou T. (2003). Introducing plants for bee-keeping at any cost? Assessment of Phacelia
tanacetifolia as nectar source plant under xeric Mediterranean conditions. Plant Syst. Evol.,
238, 155-168.
Petchey O.L., McPhearson P.T., Casey T.M. & Morin P.J. (1999). Environmental warming alters foodweb structure and ecosystem function. Nature, 402, 69-72.
49
Peters H.A., Baur B., Bazzaz F. & Körner C. (2000). Consumption rates and food preferences of slugs
in a calcareous grassland under current and future CO2 conditions. Oecologia, 125, 72-81.
Petren K. & Case T.J. (1996). An experimental demonstration of exploitation competition in an
ongoing invasion. Ecology, 77, 118-132.
Pfender W.F. & Vollmer S.S. (1999). Freezing temperature effect on survival of Puccinia graminis
subsp. graminicola in Festuca arundinacea and Lolium perenne. Plant Dis., 83, 1058-1062.
Phillips B.L. & Shine R. (2006). An invasive species induces rapid adaptive change in a native
predator: cane toads and black snakes in Australia. Proc. R. Soc. Lond. B, 273, 1545-1550.
Phillips R.L., Zak D.R., Holmes W.E. & White D.C. (2002). Microbial community composition and
function beneath temperate trees exposed to elevated atmospheric carbon dioxide and ozone.
Oecologia, 131, 236-244.
Phoenix G.K. & Press M.C. (2005). Effects of climate change on parasitic plants: The root
hemiparasitic Orobanchaceae. Folia Geobot., 40, 205-216.
Polis G.A., Hurd S.D., Jackson C.T. & Pinero F.S. (1997). El Niño effects on the dynamics and control
of an island ecosystem in the Gulf of California. Ecology, 78, 1884-1897.
Polis G.A., Sears A.L.W., Huxel G.R., Strong D.R. & Maron J. (2000). When is a trophic cascade a
trophic cascade? Trends Ecol. Evol., 15, 473-475.
Poorter H. & Navas M.L. (2003). Plant growth and competition at elevated CO2: on winners, losers and
functional groups. New Phytol., 157, 175-198.
Poulin R. & Mouritsen K.N. (2006). Climate change, parasitism and the structure of intertidal
ecosystems. J. Helminthol., 80, 183-191.
Pounds J.A., Bustamante M.R., Coloma L.A., Consuegra J.A., Fogden M.P.L., Foster P.N., La Marca
E., Masters K.L., Merino-Viteri A., Puschendorf R., Ron S.R., Sanchez-Azofeifa G.A., Still
C.J. & Young B.E. (2006). Widespread amphibian extinctions from epidemic disease driven
by global warming. Nature, 439, 161-167.
Power S.A., Ashmore M.R., Cousins D.A. & Sheppard L.J. (1998). Effects of nitrogen addition on the
stress sensitivity of Calluna vulgaris. New Phytol., 138, 663-673.
Power S.A., Green E.R., Barker C.G., Bell J.N.B. & Ashmore M.R. (2006). Ecosystem recovery:
heathland response to a reduction in nitrogen deposition. Glob. Change Biol., 12, 1241-1252.
Prado P.I. & Lewinsohn T.M. (2004). Compartments in insect-plant associations and their
consequences for community structure. J. Anim. Ecol., 73, 1168-1178.
Prange S., Gehrt S.D. & Wiggers E.P. (2003). Demographic factors contributing to high raccoon
densities in urban landscapes. J. Wildlife Manag., 67, 324-333.
Preisser E.L. & Strong D.R. (2004). Climate affects predator control of an herbivore outbreak. Am.
Nat., 163, 754-762.
Price M.V. & Waser N.M. (1998). Effects of experimental warming on plant reproductive phenology in
a subalpine meadow. Ecology, 79, 1261-1271.
Pritchard J., Griffiths B. & Hunt E.J. (2007). Can the plant-mediated impacts on aphids of elevated
CO2 and drought be predicted? Glob. Change Biol., 13, 1616-1629.
Pritekel C., Whittemore-Olson A., Snow N. & Moore J.C. (2006). Impacts from invasive plant species
and their control on the plant community and belowground ecosystem at Rocky Mountain
National Park, USA. Appl. Soil Ecol., 32, 132-141.
Prudic K.L., Oliver J.C. & Bowers M.D. (2005). Soil nutrient effects on oviposition preference, larval
performance, and chemical defense of a specialist insect herbivore. Oecologia, 143, 578-587.
Quesada M., Stoner K.E., Lobo J.A., Herrerias-Diego Y., Palacios-Guevara C., Munguia-Rosas M.A.,
Salazar K.A.O. & Rosas-Guerrero V. (2004). Effects of forest fragmentation on pollinator
activity and consequences for plant reproductive success and mating patterns in bat-pollinated
bombacaceous trees. Biotropica, 36, 131-138.
Quesada M., Stoner K.E., Rosas-Guerrero V., Palacios-Guevara C. & Lobo J.A. (2003). Effects of
habitat disruption on the activity of nectarivorous bats (Chiroptera : Phyllostomidae) in a dry
tropical forest: implications for the reproductive success of the neotropical tree Ceiba
grandiflora. Oecologia, 135, 400-406.
Rand T.A. & Louda S.M. (2004). Exotic weed invasion increases the susceptibility of native plants to
attack by a biocontrol herbivore. Ecology, 85, 1548-1554.
Rand T.A. & Tscharntke T. (2007). Contrasting effects of natural habitat loss on generalist and
specialist aphid natural enemies. Oikos, 116, 1353-1362.
Rand T.A., Tylianakis J.M. & Tscharntke T. (2006). Spillover edge effects: the dispersal of
agriculturally subsidized insect natural enemies into adjacent natural habitats. Ecol. Lett., 9,
603-614.
50
Rantalainen M.L., Fritze H., Haimi J., Pennanen T. & Setala H. (2005). Species richness and food web
structure of soil decomposer community as affected by the size of habitat fragment and habitat
corridors. Glob. Change Biol., 11, 1614-1627.
Rantalainen M.L., Haimi J., Fritze H. & Setala H. (2006). Effects of small-scale habitat fragmentation,
habitat corridors and mainland dispersal on soil decomposer organisms. Appl. Soil Ecol., 34,
152-159.
Rathcke B.J. & Jules E.S. (1993). Habitat fragmentation and plant pollinator interactions. Curr. Sci.,
65, 273-277.
Reich P.B., Tilman D., Craine J., Ellsworth D., Tjoelker M.G., Knops J., Wedin D., Naeem S.,
Bahauddin D., Goth J., Bengtson W. & Lee T.D. (2001). Do species and functional groups
differ in acquisition and use of C, N and water under varying atmospheric CO 2 and N
availability regimes? A field test with 16 grassland species. New Phytol., 150, 435-448.
Reinhart K.O. & Callaway R.M. (2006). Soil biota and invasive plants. New Phytol., 170, 445-457.
Rey A., Barton C. & Jarvis P. (1997). Belowground responses to increased atmospheric CO2
concentration in birch (Betula pendula Roth). In: Impacts of Global Change on Tree
Physiology and Forest Ecosystems. Proceedings of the International Conference on Impacts of
Global Change on Tree Physiology and Forest Ecosystems, Held 26-29 November 1996,
Wageningen (eds. Mohren G, Kramer K & Sabate S). Kluwer The Netherlands, pp. 207-212.
Richardson S.J., Press M.C., Parsons A.N. & Hartley S.E. (2002). How do nutrients and warming
impact on plant communities and their insect herbivores? A 9-year study from a sub-Arctic
heath. J. Ecol., 90, 544-556.
Rickey M.A. & Anderson R.C. (2004). Effects of nitrogen addition on the invasive grass Phragmites
australis and a native competitor Spartina pectinata. J. Appl. Ecol., 41, 888-896.
Riera N., Traveset A. & García O. (2002). Breakage of mutualisms by exotic species: the case of
Cneorum tricoccon L. in the Balearic Islands (Western Mediterranean Sea). J. Biogeogr., 29,
713-719.
Riley S.P.D., Foley J. & Chomel B. (2004). Exposure to feline and canine pathogens in bobcats and
gray foxes in urban and rural zones of a National Park in California. J. Wildl. Dis., 40, 11-22.
Rillig M.C. & Allen M.F. (1998). Arbuscular mycorrhizae of Gutierrezia sarothrae and elevated
carbon dioxide: Evidence for shifts in C allocation to and within the mycobiont. Soil Biol.
Biochem., 30, 2001-2008.
Rillig M.C. & Field C.B. (2003). Arbuscular mycorrhizae respond to plants exposed to elevated
atmospheric CO2 as a function of soil depth. Plant Soil, 254, 383-391.
Rillig M.C., Field C.B. & Allen M.F. (1999). Fungal root colonization responses in natural grasslands
after long-term exposure to elevated atmospheric CO2. Glob. Change Biol., 5, 577-585.
Rillig M.C., Hernandez G.Y. & Newton P.C.D. (2000). Arbuscular mycorrhizae respond to elevated
atmospheric CO2 after long-term exposure: evidence from a CO2 spring in New Zealand
supports the resource balance model. Ecol. Lett., 3, 475-478.
Rinnan R., Michelsen A., Baath E. & Jonasson S. (2007). Fifteen years of climate change
manipulations alter soil microbial communities in a subarctic heath ecosystem. Glob. Change
Biol., 13, 28-39.
Ris N., Allemand R., Fouillet P. & Fleury F. (2004). The joint effect of temperature and host species
induce complex genotype-by-environment interactions in the larval parasitoid of Drosophila,
Leptopilina heterotoma (Hymenoptera: Figitidae). Oikos, 106, 451-456.
Ritchie M.E. (2000). Nitrogen limitation and trophic vs. abiotic influences on insect herbivores in a
temperate grassland. Ecology, 81, 1601-1612.
Rocha O.J. & Aguilar G. (2001). Reproductive biology of the dry forest tree Enterolobium
cyclocarpum (Guanacaste) in Costa Rica: A comparison between trees left in pastures and
trees in continuous forest. Am. J. Bot., 88, 1607-1614.
Roemer G.W., Coonan T.J., Garcelon D.K., Bascompte J. & Laughrin L. (2001). Feral pigs facilitate
hyperpredation by golden eagles and indirectly cause the decline of the island fox. Anim.
Cons., 4, 307-318.
Roemer G.W., Donlan C.J. & Courchamp F. (2002). Golden eagles, feral pigs, and insular carnivores:
How exotic species turn native predators into prey. Proc. Natl Acad. Sci. U.S.A., 99, 791-796.
Roland J. & Taylor P.D. (1997). Insect parasitoid species respond to forest structure at different spatial
scales. Nature, 386, 710-713.
Root R.B. (1973). Organization of a plant-arthropod association in simple and diverse habitats: the
fauna of collards (Brassica oleracea). Ecol. Monogr., 43, 95-124.
Root T.L., Price J.T., Hall K.R., Schneider S.H., Rosenzweig C. & Pounds J.A. (2003). Fingerprints of
global warming on wild animals and plants. Nature, 421, 57-60.
51
Roschewitz I., Hucker M., Tscharntke T. & Thies C. (2005). The influence of landscape context and
farming practices on parasitism of cereal aphids. Agric. Ecosyst. Environ., 108, 218-227.
Rose K.E., Louda S.M. & Rees M. (2005). Demographic and evolutionary impacts of native and
invasive insect herbivores on Cirsium canescens. Ecology, 86, 453-465.
Rossetto M., Gross C.L., Jones R. & Hunter J. (2004). The impact of clonality on an endangered tree
(Elaeocarpus williamsianus) in a fragmented rainforest. Biol. Cons., 117, 33-39.
Rouhier H. & Read D.J. (1998). The role of mycorrhiza in determining the response of Plantago
lanceolata to CO2 enrichment. New Phytol., 139, 367-373.
Rouhier H. & Read D.J. (1999). Plant and fungal responses to elevated atmospheric CO 2 in
mycorrhizal seedlings of Betula pendula. Environ. Exp. Bot., 42, 231-241.
Rowe E.C., Healey J.R., Edwards-Jones G., Hills J., Howells M. & Jones D.L. (2006). Fertilizer
application during primary succession changes the structure of plant and herbivore
communities. Biol. Cons., 131, 510-522.
Rowles A.D. & O'Dowd D.J. (2007). Interference competition by Argentine ants displaces native ants:
implications for biotic resistance to invasion. Biol. Inv., 9, 73-85.
Roy B.A., Gusewell S. & Harte J. (2004). Response of plant pathogens and herbivores to a warming
experiment. Ecology, 85, 2570-2581.
Rudgers J.A., Holah J., Orr S.P. & Clay K. (2007). Forest succession suppressed by an introduced
plant-fungal symbiosis. Ecology, 88, 18-25.
Ruess L., Michelsen A., Schmidt I.K. & Jonasson S. (1999). Simulated climate change affecting
microorganisms, nematode density and biodiversity in subarctic soils. Plant Soil, 212, 63-73.
Russell F.L. & Louda S.M. (2004). Phenological synchrony affects interaction strength of an exotic
weevil with Platte thistle, a native host plant. Oecologia, 139, 525-534.
Russell F.L. & Louda S.M. (2005). Indirect interaction between two native thistles mediated by an
invasive exotic floral herbivore. Oecologia, 146, 373-384.
Rusterholz H.P. & Erhardt A. (1998). Effects of elevated CO2 on flowering phenology and nectar
production of nectar plants important for butterflies of calcareous grasslands. Oecologia, 113,
341-349.
Ryall K.L. & Fahrig L. (2005). Habitat loss decreases predator-prey ratios in a pine-bark beetle system.
Oikos, 110, 265-270.
Ryall K.L. & Fahrig L. (2006). Response of predators to loss and fragmentation of prey habitat: A
review of theory. Ecology, 87, 1086-1093.
Saavedra F., Inouye D.W., Price M.V. & Harte J. (2003). Changes in flowering and abundance of
Delphinium nuttallianum (Ranunculaceae) in response to a subalpine climate warming
experiment. Glob. Change Biol., 9, 885-894.
Saggar S., McIntosh P.D., Hedley C.B. & Knicker H. (1999). Changes in soil microbial biomass,
metabolic quotient, and organic matter turnover under Hieracium (H. pilosella L.). Biol. Fert.
Soils, 30, 232-238.
Salminen S.O., Richmond D.S., Grewal S.K. & Grewal P.S. (2005). Influence of temperature on
alkaloid levels and fall armyworm performance in endophytic tall fescue and perennial
ryegrass. Ent. Exp. Appl., 115, 417-426.
Sanders I.R., Streitwolf-Engel R., van der Heijden M.G.A., Boller T. & Wiemken A. (1998). Increased
allocation to external hyphae of arbuscular mycorrhizal fungi under CO2 enrichment.
Oecologia, 117, 496-503.
Sanders N.J., Belote R.T. & Weltzin J.F. (2004). Multitrophic effects of elevated atmospheric CO2 on
understory plant and arthropod communities. Environ. Ent., 33, 1609-1616.
Schauber E.M., Kelly D., Turchin P., Simon C., Lee W.G., Allen R.B., Payton I.J., Wilson P.R.,
Cowan P.E. & Brockie R.E. (2002). Masting by eighteen New Zealand plant species: The role
of temperature as a synchronizing cue. Ecology, 83, 1214-1225.
Scheu S. & Schaefer M. (1998). Bottom-up control of the soil macrofauna community in a beechwood
on limestone: Manipulation of food resources. Ecology, 79, 1573-1585.
Schierenbeck K.A., Mack R.N. & Sharitz R.R. (1994). Effects of herbivory on growth and biomass
allocation in native and introduced species of Lonicera. Ecology, 75, 1661-1672.
Schiffman P.M. (1994). Promotion of exotic weed establishment by endangered giant kangaroo rats
(Dipodomys ingens) in a California grassland. Biodiv. Conserv., 3, 524-537.
Schmidt K. & Jensen K. (2000). Genetic structure and AFLP variation of remnant populations in the
rare plant Pedicularis palustris (Scrophulariaceae) and its relation to population size and
reproductive components. Am. J. Bot., 87, 678-689.
52
Schmidt O., Curry J.P., Hackett R.A., Purvis G. & Clements R.O. (2001). Earthworm communities in
conventional wheat monocropping and low-input wheat-clover intercropping systems. Ann.
Appl. Biol., 138, 377-388.
Schneider K., Scheu S. & Maraun M. (2007). Microarthropod density and diversity respond little to
spatial isolation. Basic Appl. Ecol., 8, 26-35.
Schwartz M.W., Hoeksema J.D., Gehring C.A., Johnson N.C., Klironomos J.N., Abbott L.K. & Pringle
A. (2006). The promise and the potential consequences of the global transport of mycorrhizal
fungal inoculum. Ecol. Lett., 9, 501-515.
Sessions L. & Kelly D. (2002). Predator-mediated apparent competition between an introduced grass,
Agrostis capillaris, and a native fern, Botrychium australe (Ophioglossaceae), in New
Zealand. Oikos, 96, 102-109.
Settle W.H. & Wilson L.T. (1990). Invasion by the variegated leafhopper and biotic interactions parasitism, competition, and apparent competition. Ecology, 71, 1461-1470.
Severns P. (2003). Inbreeding and small population size reduce seed set in a threatened and fragmented
plant species, Lupinus sulphureus spp. kincaidii (Fabaceae). Biol. Cons., 110, 221-229.
Siguenza C., Crowley D.E. & Allen E.B. (2006). Soil microorganisms of a native shrub and exotic
grasses along a nitrogen deposition gradient in southern California. Appl. Soil Ecol., 32, 1326.
Siitonen P., Lehtinen A. & Siitonen M. (2005). Effects of forest edges on the distribution, abundance,
and regional persistence of wood-rotting fungi. Cons. Biol., 19, 250-260.
Silliman B.R. & Bertness M.D. (2004). Shoreline development drives invasion of Phragmites australis
and the loss of plant diversity on New England salt marshes. Cons. Biol., 18, 1424-1434.
Skirvin D.J., Perry J.N. & Harrington R. (1997). The effect of climate change on an aphid-coccinellid
interaction. Glob. Change Biol., 3, 1-11.
Smith-Ramirez C. & Armesto J.J. (2003). Foraging behaviour of bird pollinators on Embothrium
coccineum (Proteaceae) trees in forest fragments and pastures in southern Chile. Austral Ecol.,
28, 53-60.
Smith K.F. & Carpenter S.M. (2006). Potential spread of introduced black rat (Rattus rattus) parasites
to endemic deer mice (Peromyscus maniculatus) on the California Channel Islands. Divers.
Distrib., 12, 742-748.
Snyder W.E. & Evans E.W. (2006). Ecological effects of invasive arthropod generalist predators. Ann.
Rev. Ecol. Evol. Syst., 37, 95-122.
Soderstrom B., Baath E. & Lundgren B. (1983). Decrease in soil microbial activity and biomasses
owing to nitrogen amendments. Can. J. Microbiol., 29, 1500-1506.
Sohlenius B. & Boström S. (1999). Effects of climate change on soil factors and metazoan microfauna
(nematodes, tardigrades and rotifers) in a Swedish tundra soil - a soil transplantation
experiment. Appl. Soil Ecol., 12, 113-128.
Somanathan H. & Borges R.M. (2000). Influence of exploitation on population structure, spatial
distribution and reproductive success of dioecious species in a fragmented cloud forest in
India. Biol. Cons., 94, 243-256.
Sonnemann I. & Wolters V. (2005). The microfood web of grassland soils responds to a moderate
increase in atmospheric CO2. Glob. Change Biol., 11, 1148-1155.
Sparks T.H. & Yates T.J. (1997). The effect of spring temperature on the appearance dates of British
butterflies 1883-1993. Ecography, 20, 368-374.
Spiegelberger T., Matthies D., Muller-Scharer H. & Schaffner U. (2006). Scale-dependent effects of
land use on plant species richness of mountain grassland in the European Alps. Ecography,
29, 541-548.
Spurr E.B. & Anderson S.H. (2004). Bird species diversity and abundance before and after eradication
of possums and wallabies on Rangitoto Island, Hauraki Gulf, New Zealand. N. Z. J. Ecol., 28,
143-149.
Stacey D.A. (2003). Climate and biological control in organic crops. Intl J. Pest Manag., 49, 205-214.
Stacey D.A. & Fellowes M.D.E. (2002). Influence of elevated CO2 on interspecific interactions at
higher trophic levels. Glob. Change Biol., 8, 668-678.
Staddon P.L., Fitter A.H. & Graves J.D. (1999). Effect of elevated atmospheric CO2 on mycorrhizal
colonization, external mycorrhizal hyphal production and phosphorus inflow in Plantago
lanceolata and Trifolium repens in association with the arbuscular mycorrhizal fungus Glomus
mosseae. Glob. Change Biol., 5, 347-358.
Staddon P.L., Graves J.D. & Fitter A.H. (1998). Effect of enhanced atmospheric CO2 on mycorrhizal
colonization by Glomus mosseae in Plantago lanceolata and Trifolium repens. New Phytol.,
139, 571-580.
53
Staddon P.L., Gregersen R. & Jakobsen I. (2004). The response of two Glomus mycorrhizal fungi and a
fine endophyte to elevated atmospheric CO2, soil warming and drought. Glob. Change Biol.,
10, 1909-1921.
Staddon P.L., Heinemeyer A. & Fitter A.H. (2002). Mycorrhizas and global environmental change:
research at different scales. Plant Soil, 244, 253-261.
Stahl J., Van Der Graaf A.J., Drent R.H. & Bakker J.P. (2006). Subtle interplay of competition and
facilitation among small herbivores in coastal grasslands. Funct. Ecol., 20, 908-915.
Standish R.J. (2004). Impact of an invasive clonal herb on epigaeic invertebrates in forest remnants in
New Zealand. Biol. Cons., 116, 49-58.
Stapp P., Antolin M.F. & Ball M. (2004). Patterns of extinction in prairie dog metapopulations: plague
outbreaks follow El Niño events. Front. Ecol. Environ., 2, 235-240.
Steffan-Dewenter I., Münzenberg U., Bürger C., Thies C. & Tscharntke T. (2002). Scale-dependent
effects of landscape context on three pollinator guilds. Ecology, 83, 1421-1432.
Steffan-Dewenter I., Münzenberg U. & Tscharntke T. (2001). Pollination, seed set and seed predation
on a landscape scale. Proc.R.Soc.Lond.B, 268, 1685-1690.
Steffan-Dewenter I. & Tscharntke T. (1999). Effects of habitat isolation on pollinator communities and
seed set. Oecologia, 121, 432-440.
Stevens C.J., Dise N.B., Mountford J.O. & Gowing D.J. (2004). Impact of nitrogen deposition on the
species richness of grasslands. Science, 303, 1876-1879.
Stevens G.N. & Jones R.H. (2006). Patterns in soil fertility and root herbivory interact to influence
fine-root dynamics. Ecology, 87, 616-624.
Sticht C., Schrader S., Giesemann A. & Weigel H.J. (2006). Effects of elevated atmospheric CO2 and
N fertilization on abundance, diversity and C-isotopic signature of collembolan communities
in arable soil. Appl. Soil Ecol., 34, 219-229.
Stiling P., Cattell M., Moon D.C., Rossi A., Hungate B.A., Hymus G. & Drake B. (2002). Elevated
atmospheric CO2 lowers herbivore abundance, but increases leaf abscission rates. Glob.
Change Biol., 8, 658-667.
Stiling P. & Cornelissen T. (2007). How does elevated carbon dioxide (CO2) affect plant-herbivore
interactions? A field experiment and meta-analysis of CO2-mediated changes on plant
chemistry and herbivore performance. Glob. Change Biol., 13, 1823-1842.
Stiling P., Moon D., Hymus G. & Drake B. (2004). Differential effects of elevated CO2 on acorn
density, weight, germination, and predation among three oak species in a scrub-oak forest.
Glob. Change Biol., 10, 228-232.
Stiling P. & Moon D.C. (2005). Quality or quantity: the direct and indirect effects of host plants on
herbivores and their natural enemies. Oecologia, 142, 413-420.
Stiling P., Rossi A.M., Hungate B., Dijkstra P., Hinkle C.R., Knott W.M. & Drake B. (1999).
Decreased leaf-miner abundance in elevated CO2: Reduced leaf quality and increased
parasitoid attack. Ecol. Appl., 9, 240-244.
Stinson K.A., Campbell S.A., Powell J.R., Wolfe B.E., Callaway R.M., Thelen G.C., Hallett S.G., Prati
D. & Klironomos J.N. (2006). Invasive plant suppresses the growth of native tree seedlings by
disrupting belowground mutualisms. PLoS Biol., 4, 727-731.
Stireman J.O., Dyer L.A., Janzen D.H., Singer M.S., Li J.T., Marquis R.J., Ricklefs R.E., Gentry G.L.,
Hallwachs W., Coley P.D., Barone J.A., Greeney H.F., Connahs H., Barbosa P., Morais H.C.
& Diniz I.R. (2005). Climatic unpredictability and parasitism of caterpillars: Implications of
global warming. Proc. Natl Acad. Sci. U.S.A., 102, 17384-17387.
Stoll P., Dolt C., Goverde M. & Baur B. (2006). Experimental habitat fragmentation and invertebrate
grazing in a herbaceous grassland species. Basic Appl. Ecol., 7, 307-319.
Stout J.C., Kells A.R. & Goulson D. (2002). Pollination of the invasive exotic shrub Lupinus arboreus
(Fabaceae) by introduced bees in Tasmania. Biol. Cons., 106, 425-434.
Strayer D.L., Eviner V.T., Jeschke J.M. & Pace M.L. (2006). Understanding the long-term effects of
species invasions. Trends Ecol. Evol., 21, 645-651.
Strengbom J., Englund G. & Ericson L. (2006). Experimental scale and precipitation modify effects of
nitrogen addition on a plant pathogen. J. Ecol., 94, 227-233.
Strengbom J., Nordin A., Nasholm T. & Ericson L. (2002). Parasitic fungus mediates change in
nitrogen-exposed boreal forest vegetation. J. Ecol., 90, 61-67.
Stromberger M., Shah Z. & Westfall D. (2007). Soil microbial communities of no-till dryland
agroecosystems across an evapotranspiration gradient. Appl. Soil Ecol., 35, 94-106.
Suarez A.V. & Case T.J. (2002). Bottom-up effects on persistence of a specialist predator: Ant
invasions and horned lizards. Ecol. Appl., 12, 291-298.
54
Sudderth E.A., Stinson K.A. & Bazzaz F.A. (2005). Host-specific aphid population responses to
elevated CO2 and increased N availability. Glob. Change Biol., 11, 1997-2008.
Suttle K.B. & Hoddle M.S. (2006). Engineering enemy-free space: an invasive pest that kills its
predators. Biol. Inv., 8, 639-649.
Swihart R.K., Feng Z., Slade N.A., Mason D.M. & Gehring T.M. (2001). Effects of habitat destruction
and resource supplementation in a predator-prey metapopulation model. J. Theor. Biol., 210,
287-303.
Tallmon D.A., Jules E.S., Radke N.J. & Mills L.S. (2003). Of mice and men and trillium: Cascading
effects of forest fragmentation. Ecol. Appl., 13, 1193-1203.
Telfer S., Bown K.J., Sekules R., Begon I., Hayden T. & Birtles R. (2005). Disruption of a hostparasite system following the introduction of an exotic host species. Parasitology, 130, 661668.
Tewksbury J., Levey D., Haddad N., Sargent S., Orrock J., Weldon A., Danielson B., Brinkerhoff J.,
Damschen E. & Townsend P. (2002). Corridors affect plants, animals, and their interactions in
fragmented landscapes. Proc. Natl Acad. Sci. U.S.A., 99, 12923-12926.
Tewksbury J.J., Garner L., Garner S., Lloyd J.D., Saab V. & Martin T.E. (2006). Tests of landscape
influence: Nest predation and brood parasitism in fragmented ecosystems. Ecology, 87, 759768.
Thies C. & Tscharntke T. (1999). Landscape structure and biological control in agroecosystems.
Science, 285, 893-895.
Thompson C.M. & Gese E.M. (2007). Food webs and intraguild predation: community interactions of a
native mesocarnivore. Ecology, 88, 334-346.
Thompson J.D. (2001). How do visitation patterns vary among pollinators in relation to floral display
and floral design in a generalist pollination system? Oecologia, 126, 386-394.
Throop H.L. (2005). Nitrogen deposition and herbivory affect biomass production and allocation in an
annual plant. Oikos, 111, 91-100.
Throop H.L. & Lerdau M.T. (2004). Effects of nitrogen deposition on insect herbivory: Implications
for community and ecosystem processes. Ecosyst., 7, 109-133.
Tilman D. & Lehman C.L. (2001). Human-caused environmental change: impacts on plant diversity
and evolution. Proc. Natl Acad. Sci. U.S.A., 98, 5433-5440.
Tomimatsu H. & Ohara M. (2002). Effects of forest fragmentation on seed production of the
understory herb Trillium camschatcense. Cons. Biol., 16, 1277-1285.
Tompkins D.M. & Gleeson D.M. (2006). Relationship between avian malaria distribution and an exotic
invasive mosquito in New Zealand. J. R. Soc. N.Z., 36, 51-62.
Tompkins D.M., Sainsbury A.W., Nettleton P., Buxton D. & Gurnell J. (2002). Parapoxvirus causes a
deleterious disease in red squirrels associated with UK population declines. Proc. R. Soc.
Lond. B, 269, 529-533.
Tompkins D.M., White A.R. & Boots M. (2003). Ecological replacement of native red squirrels by
invasive greys driven by disease. Ecol. Lett., 6, 189-196.
Toussaint J.F., Kerkhofs P. & De Clercq K. (2006). Influence of global climate changes on arboviruses
spread. Ann. Med. Vet., 150, 56-63.
Traveset A. & Richardson D.M. (2006). Biological invasions as disruptors of plant reproductive
mutualisms. Trends Ecol. Evol., 21, 208-216.
Traveset A. & Riera N. (2005). Disruption of a plant-lizard seed dispersal system and its ecological
effects on a threatened endemic plant in the Balearic Islands. Cons. Biol., 19, 421-431.
Traveset A. & Saez E. (1997). Pollination of Euphorbia dendroides by lizards and insects: Spatiotemporal variation in patterns of flower visitation. Oecologia, 111, 241-248.
Treseder K.K. (2004). A meta-analysis of mycorrhizal responses to nitrogen, phosphorus, and
atmospheric CO2 in field studies. New Phytol., 164, 347-355.
Treseder K.K. & Vitousek P.M. (2001). Effects of soil nutrient availability on investment in acquisition
of N and P in Hawaiian rain forests. Ecology, 82, 946-954.
Tscharntke T. & Brandl R. (2004). Plant-insect interactions in fragmented landscapes. Ann. Rev. Ent.,
49, 405-430.
Turnock W.J., Wise I.L. & Matheson F.O. (2003). Abundance of some native coccinellines
(Coleoptera: Coccinellidae) before and after the appearance of Coccinella septempunctata.
Can. Entomol., 135, 391-404.
Tylianakis J.M., Klein A.M., Lozada T. & Tscharntke T. (2006). Spatial scale of observation affects
alpha, beta and gamma diversity of cavity-nesting bees and wasps across a tropical land-use
gradient. J. Biogeogr., 33, 1295-1304.
55
Tylianakis J.M., Klein A.M. & Tscharntke T. (2005). Spatiotemporal variation in the diversity of
hymenoptera across a tropical habitat gradient. Ecology, 86, 3296-3302.
Tylianakis J.M., Tscharntke T. & Lewis O.T. (2007). Habitat modification alters the structure of
tropical host-parasitoid food webs. Nature, 445, 202-205.
Valdivia C.E. & Simonetti J.A. (2007). Decreased frugivory and seed germination rate do not reduce
seedling recruitment rates of Aristotelia chilensis in a fragmented forest. Biodiv. Conserv., 16,
1593-1602.
Valdivia C.E., Simonetti J.A. & Henriquez C.A. (2006). Depressed pollination of Lapageria rosea
Ruiz et pav. (Philesiaceae) in the fragmented temperate rainforest of Southern South America.
Biodiv. Conserv., 15, 1845-1856.
Valentine L.E., Roberts B. & Schwarzkopf L. (2007). Mechanisms driving avoidance of non-native
plants by lizards. J. Appl. Ecol., 44, 228-237.
Valladares G., Salvo A. & Cagnolo L. (2006). Habitat fragmentation effects on trophic processes of
insect-plant food webs. Cons. Biol., 20, 212-217.
Van der Putten W.H., Klironomos J.N. & Wardle D.A. (2007). Microbial ecology of biological
invasions. The ISME Journal, 1, 28-37.
van Nouhuys S. (2005). Effects of habitat fragmentation at different trophic levels in insect
communities. Ann. Zool. Fennici, 42, 433-447.
Van Nouhuys S. & Lei G.C. (2004). Parasitoid-host metapopulation dynamics: the causes and
consequences of phenological asynchrony. J. Anim. Ecol., 73, 526-535.
van Rossum F., Echchgadda G., Szabadi I. & Triest L. (2002). Commonness and long-term survivial in
fragmented habitats: Primula elatior as a study case. Cons. Biol., 16, 1286-1295.
Vazquez D.P. & Simberloff D. (2004). Indirect effects of an introduced ungulate on pollination and
plant reproduction. Ecol. Monogr., 74, 281-308.
Veteli T.O., Kuokkanen K., Julkunen-Tiitto R., Roininen H. & Tahvanainen J. (2002). Effects of
elevated CO2 and temperature on plant growth and herbivore defensive chemistry. Glob.
Change Biol., 8, 1240-1252.
Virtanen T. & Neuvonen S. (1999). Performance of moth larvae on birch in relation to altitude, climate,
host quality and parasitoids. Oecologia, 120, 92-101.
Visser M.E., Adriaensen F., van Balen J.H., Blondel J., Dhondt A.A., van Dongen S., du Feu C.,
Ivankina E.V., Kerimov A.B., de Laet J., Matthysen E., McCleery R., Orell M. & Thomson
D.L. (2003). Variable responses to large-scale climate change in European Parus populations.
Proc. R. Soc. Lond. B, 270, 367-372.
Visser M.E. & Both C. (2005). Shifts in phenology due to global climate change: the need for a
yardstick. Proc. R. Soc. Lond. B, 272, 2561-2569.
Visser M.E. & Holleman L.J.M. (2001). Warmer springs disrupt the synchrony of oak and winter moth
phenology. Proc. R. Soc. Lond. B, 268, 289-294.
Voigt W., Perner J., Davis A.J., Eggers T., Schumacher J., Bärhmann R., Fabian B., Heinrich W.,
Köhler G., Lichter D., Marstaller R. & Sander F.W. (2003). Trophic levels are differentially
sensitive to climate. Ecology, 84, 2444-2453.
Vuorinen T., Nerg A.M., Ibrahim M.A., Reddy G.V.P. & Holopainen J.K. (2004). Emission of Plutella
xylostella-induced compounds from cabbages grown at elevated CO 2 and orientation behavior
of the natural enemies. Plant Physiology, 135, 1984-1992.
Wagenius S., Lonsdorf E. & Neuhauser C. (2007). Patch aging and the S-allee effect: Breeding system
effects on the demographic response of plants to habitat fragmentation. Am. Nat., 169, 383397.
Waldrop M.P. & Firestone M.K. (2006). Response of microbial community composition and function
to soil climate change. Microb. Ecol., 52, 716-724.
Walker M.D., Wahren C.H., Hollister R.D., Henry G.H.R., Ahlquist L.E., Alatalo J.M., Bret-Harte
M.S., Calef M.P., Callaghan T.V., Carroll A.B., Epstein H.E., Jonsdottir I.S., Klein J.A.,
Magnusson B., Molau U., Oberbauer S.F., Rewa S.P., Robinson C.H., Shaver G.R., Suding
K.N., Thompson C.C., Tolvanen A., Totland O., Turner P.L., Tweedie C.E., Webber P.J. &
Wookey P.A. (2006). Plant community responses to experimental warming across the tundra
biome. Proc. Natl Acad. Sci. U.S.A., 103, 1342-1346.
Wall M.A., Timmerman-Erskine M. & Boyd R.S. (2003). Conservation impact of climatic variability
on pollination of the federally endangered plant, Clematis socialis (Ranunculaceae).
Southeast. Nat., 2, 11-24.
Wang Q., Wang C.H., Zhao B., Ma Z.J., Luo Y.Q., Chen J.K. & Li B. (2006). Effects of growing
conditions on the growth of and interactions between salt marsh plants: Implications for
invasibility of habitats. Biol. Inv., 8, 1547-1560.
56
Warburton C.L., James E.A., Fripp Y.J., Trueman S.J. & Wallace H.M. (2000). Clonality and sexual
reproductive failure in remnant populations of Santalum lanceolatum (Santalaceae). Biol.
Cons., 96, 45-54.
Ward M. & Johnson S.D. (2005). Pollen limitation and demographic structure in small fragmented
populations of Brunsvigia radulosa (Amaryllidaceae). Oikos, 108, 253-262.
Wardle D.A. (1995). Impact of disturbance on detritus food-webs in agro-ecosystems of contrasting
tillage and weed management practices. Adv. Ecol. Res., 26, 105-185.
Wardle D.A. (2002). Communities and Ecosystems: Linking the Aboveground and Belowground
Components. Princeton University Press, Princeton USA.
Wardle D.A., Barker G.M., Yeates G.W., Bonner K.I. & Ghani A. (2001). Introduced browsing
mammals in New Zealand natural forests: aboveground and belowground consequences. Ecol.
Monogr., 71, 587-614.
Wardle D.A., Yeates G.W., Nicholson K.S., Bonner K.I. & Watson R.N. (1999). Response of soil
microbial biomass dynamics, activity and plant litter decomposition to agricultural
intensification over a seven-year period. Soil Biol. Biochem., 31, 1707-1720.
Wasley J., Robinson S.A., Lovelock C.E. & Popp M. (2006). Climate change manipulations show
Antarctic flora is more strongly affected by elevated nutrients than water. Glob. Change Biol.,
12, 1800-1812.
Watling J.R. & Press M.C. (1997). How is the relationship between the C-4 cereal Sorghum bicolor
and the C-3 root hemi-parasites Striga hermonthica and Striga asiatica affected by elevated
CO2? Plant Cell Environ., 20, 1292-1300.
Watling J.R. & Press M.C. (1998). How does the C-4 grass Eragrostis pilosa respond to elevated
carbon dioxide and infection with the parasitic angiosperm Striga hermonthica? New Phytol.,
140, 667-675.
Watling J.R. & Press M.C. (2000). Infection with the parasitic angiosperm Striga hermonthica
influences the response of the C-3 cereal Oryza sativa to elevated CO2. Glob. Change Biol., 6,
919-930.
Watts C.H. & Didham R.K. (2006). Influences of habitat isolation on invertebrate colonization of
Sporadanthus ferrugineus in a mined peat bog. Restor. Ecol., 14, 412-419.
West J.B., HilleRisLambers J., Lee T.D., Hobbie S.E. & Reich P.B. (2005). Legume species identity
and soil nitrogen supply determine symbiotic nitrogen-fixation responses to elevated
atmospheric [CO2]. New Phytol., 167, 523-530.
White E.M., Wilson J.C. & Clarke A.R. (2006). Biotic indirect effects: a neglected concept in invasion
biology. Divers. Distrib., 12, 443-455.
Wiemken V., Laczko E., Ineichen K. & Boller T. (2001). Effects of elevated carbon dioxide and
nitrogen fertilization on mycorrhizal fine roots and the soil microbial community in beechspruce ecosystems on siliceous and calcareous soil. Microb. Ecol., 42, 126-135.
Williams R.S., Lincoln D.E. & Norby R.J. (2003). Development of gypsy moth larvae feeding on red
maple saplings at elevated CO2 and temperature. Oecologia, 137, 114-122.
Williams R.S., Norby R.J. & Lincoln D.E. (2000). Effects of elevated CO2 and temperature-grown red
and sugar maple on gypsy moth performance. Glob. Change Biol., 6, 685-695.
Willis A.J. & Memmott J. (2005). The potential for indirect effects between a weed, one of its
biocontrol agents and native herbivores: A food web approach. Biol. Control, 35, 299-306.
Wilson J.D., Morris A.J., Arroyo B.E., Clark S.C. & Bradbury R.B. (1999). A review of the abundance
and diversity of invertebrate and plant foods of granivorous birds in northern Europe in
relation to agricultural change. Agric. Ecosyst. Environ., 75, 13-30.
Wilson S.D. & Tilman D. (1991). Components of plant competition along an experimental gradient of
nitrogen availability. Ecology, 72, 1050-1065.
Winfree R., Griswold T. & Kremen C. (2007). Effect of human disturbance on bee communities in a
forested ecosystem. Cons. Biol., 21, 213-223.
Witt A.B.R. & Giliomee J.H. (2004). The impact of an invasive ant, Linepithema humile (Mayr), on the
dispersal of Phylica pubescens Aiton seeds in South Africa. Afr. Entomol., 12, 179-185.
Witzell J. & Shevtsova A. (2004). Nitrogen-induced changes in phenolics of Vaccinium myrtillus Implications for interaction with a parasitic fungus. J. Chem. Ecol., 30, 1937-1956.
Wolf A.T. & Harrison S.P. (2001). Effects of habitat size and patch isolation on reproductive success
of the serpentine morning glory. Cons. Biol., 15, 111-121.
Wolf J., Johnson N.C., Rowland D.L. & Reich P.B. (2003). Elevated CO2 and plant species richness
impact arbuscular mycorrhizal fungal spore communities. New Phytol., 157, 579-588.
Worthy F.R., Law R. & Hulme P.E. (2006). Modelling the quantitative effects of pre- and postdispersal seed predation in Pinus sylvestris L. J. Ecol., 94, 1201-1213.
57
Wright A.N. & Gompper M.E. (2005). Altered parasite assemblages in raccoons in response to
manipulated resource availability. Oecologia, 144, 148-156.
Wu S.M., Hu D.X. & Ingham E.R. (2005). Comparison of soil biota between organic and conventional
agroecosystems in Oregon, USA. Pedosphere, 15, 395-403.
Wu T.H., Chellemi D.O., Martin K.J., Graham J.H. & Rosskop E.N. (2007). Discriminating the effects
of agricultural land management practices on soil fungal communities. Soil Biol. Biochem.,
39, 1139-1155.
Yanoviak S.P., Paredes J.E.R., Lounibos L.P. & Weaver S.C. (2006). Deforestation alters phytotelm
habitat availability and mosquito production in the Peruvian Amazon. Ecol. Appl., 16, 18541864.
Yasuda H., Evans E.W., Kajita Y., Urakawa K. & Takizawa T. (2004). Asymmetric larval interactions
between introduced and indigenous ladybirds in North America. Oecologia, 141, 722-731.
Yates C.J. & Ladd P.G. (2005). Relative importance of reproductive biology and establishment ecology
for persistence of a rare shrub in a fragmented landscape. Cons. Biol., 19, 239-249.
Yeates G.W., Bardgett R.D., Cook R., Hobbs P.J., Bowling P.J. & Potter J.F. (1997). Faunal and
microbial diversity in three Welsh grassland soils under conventional and organic
management regimes. J. Appl. Ecol., 34, 453-470.
Yeates G.W., Newton P.C.D. & Ross D.J. (2003). Significant changes in soil microfauna in grazed
pasture under elevated carbon dioxide. Biol. Fert. Soils, 38, 319-326.
Yeates G.W., Wardle D.A. & Watson R.N. (1999). Responses of soil nematode populations,
community structure, diversity and temporal variability to agricultural intensification over a
seven-year period. Soil Biol. Biochem., 31, 1721-1733.
Yeates G.W. & Williams P.A. (2001). Influence of three invasive weeds and site factors on soil
microfauna in New Zealand. Pedobiologia, 45, 367-383.
Zak D.R., Pregitzer K.S., King J.S. & Holmes W.E. (2000). Elevated atmospheric CO2, fine roots and
the response of soil microorganisms: a review and hypothesis. New Phytol., 147, 201-222.
Zavaleta E.S., Thomas B.D., Chiariello N.R., Asner G.P., Shaw M.R. & Field C.B. (2003). Plants
reverse warming effect on ecosystem water balance. Proc. Natl Acad. Sci. U.S.A., 100, 98929893.
Zaviezo T., Grez A.A., Estades C.F. & Perez A. (2006). Effects of habitat loss, habitat fragmentation,
and isolation on the density, species richness, and distribution of ladybeetles in manipulated
alfalfa landscapes. Ecol. Ent., 31, 646-656.
Zhang W., Parker K.M., Luo Y., Wan S., Wallace L.L. & Hu S. (2005). Soil microbial responses to
experimental warming and clipping in a tallgrass prairie. Glob. Change Biol., 11, 266-277.
Zvereva E.L. & Kozlov M.V. (2006). Consequences of simultaneous elevation of carbon dioxide and
temperature for plant-herbivore interactions: a metaanalysis. Glob. Change Biol., 12, 27-41.