This study investigated the effects of habitat restoration in 12 urban parks in Metro Vancouver on plant and pollinator communities. Restored plots had higher plant species richness and diversity compared to control plots, but similar plant abundance. Pollinator abundance, richness and diversity were not significantly different between restored and control plots. Network analysis found control plots had higher asymmetry, suggesting invasive plants increase network resilience. The results suggest that while restorations improved plant diversity, added native plants did not provide enough additional floral resources to significantly change pollinator communities compared to resources from invasive species in control plots. Managers should ensure alternative forage is available after invasive removal by planting generalist native species with overlapping blooms.
Filling the Gaps: Arbuscular Mycorrhizal Fungi Biodiversity in the Tropical E...CrimsonpublishersMCDA
Filling the Gaps: Arbuscular Mycorrhizal Fungi Biodiversity in the Tropical Ecosystems by Geofrey E Soka* in Modern Concepts & Developments in Agronomy
Filling the Gaps: Arbuscular Mycorrhizal Fungi Biodiversity in the Tropical E...CrimsonpublishersMCDA
Filling the Gaps: Arbuscular Mycorrhizal Fungi Biodiversity in the Tropical Ecosystems by Geofrey E Soka* in Modern Concepts & Developments in Agronomy
Plant species and communities assessment in interaction with edaphic and topo...Shujaul Mulk Khan
The current analyses of vegetation were aimed to study the different effects of environmental variables on plant species and communities and their combined interactions to these variables, identified threats to local vegetation and suggestion for remedial measures in the Mount Eelum, Swat, Pakistan. For assessment of environmental variability quantitative ecological techniques were used through quadrats having sizes of 2 × 2, 5 × 5 and 10 × 10 m2 for herbs, shrubs and trees respectively. Result of the present study revealed 124 plant species in the study area. Canonical Correspondence Analysis (CCA) was used to analyze the ecological gradient of vegetation. The environmental data and species abundance were used in CANOCO software version 4.5. The presence absence data of plant species were elaborated with Cluster and Two Way Cluster Analysis techniques using PC-ORD version 5 to show different species composition that resulted in five plant communities. Findings indicate that elevation, aspect and soil texture are the strongest variables that have significant effect on species composition and distribution of various communities shown with P value 0.0500. It is recommended to protect and use sensibly whole of the Flora normally and rare species particularly in the region.
Genetic diversity enhances the resistance of aseagrass ecosyMatthewTennant613
Genetic diversity enhances the resistance of a
seagrass ecosystem to disturbance
A. Randall Hughes* and John J. Stachowicz
Section of Evolution and Ecology, One Shields Avenue, University of California, Davis, CA 95616
Edited by G. David Tilman, University of Minnesota, St. Paul, MN, and approved May 4, 2004 (received for review April 14, 2004)
Motivated by recent global reductions in biodiversity, empirical
and theoretical research suggests that more species-rich systems
exhibit enhanced productivity, nutrient cycling, or resistance to
disturbance or invasion relative to systems with fewer species. In
contrast, few data are available to assess the potential ecosystem-
level importance of genetic diversity within species known to play
a major functional role. Using a manipulative field experiment, we
show that increasing genotypic diversity in a habitat-forming
species (the seagrass Zostera marina) enhances community resis-
tance to disturbance by grazing geese. The time required for
recovery to near predisturbance densities also decreases with
increasing eelgrass genotypic diversity. However, there is no effect
of diversity on resilience, measured as the rate of shoot recovery
after the disturbance, suggesting that more rapid recovery in
diverse plots is due solely to differences in disturbance resistance.
Genotypic diversity did not affect ecosystem processes in the
absence of disturbance. Thus, our results suggest that genetic
diversity, like species diversity, may be most important for enhanc-
ing the consistency and reliability of ecosystems by providing
biological insurance against environmental change.
There is growing recognition that humans are highly depen-dent on natural ecosystems for a variety of goods and
services (1). Maintaining the provision of these goods and
services in the face of natural and anthropogenic disturbances is
critical to achieving both conservation and economic goals.
Motivated by accelerating rates of worldwide decline in biodi-
versity (2), considerable research has focused on the conse
quences of local species loss for goods and services provided by
ecosystems (2– 8). Much of this work focuses on the effects of
declining species richness on short-term processes such as pro-
duction, community respiration, and nutrient cycling (2). Al-
though the results are far from unequivocal and subject to
varying interpretation (e.g., ref. 9), it does appear that, in some
systems, reductions in local species diversity contribute to a
decline in ecosystem properties such as productivity and resis-
tance to disturbance (see review in ref. 2).
Nevertheless, many important ecosystems, such as kelp forests,
cattail marshes, and fir forests, are dominated by, and dependent
on, one or a few key plant species (10). Furthermore, individual
predator and herbivore species often play a disproportionate role in
determining ecosystem processes, overwhelming any effect of spe-
cies diversity (11). Dominant, numerically abundant s ...
Effect of glyphosate herbicide on the behavior of soil arthropods in non-orga...Salah Hussein
The objective of this study was to assess the effect of the glyphosate
application on the population of soil arthropods (collected with pitfall and
Berlese funnels) and their diversity after application of herbicide. Results
of soil arthropods collected with pitfall traps showed that glyphosate
herbicide has played a great role in detecting the activity of different
groups of soil arthropods with different reduction percentages. In insects
caused -23.7%, mites +48.8%, other arthropods -66.7% and total
population -23.3% reduction. Also glyphosate herbicide reduced the
diversity from 2.2 (ShanoonWiner values) to 1.6, as well as the equitability
%, from 46.6 % in pretreatment to 28.5% in post treatment. While it
caused a highly significant increase in the insect's population with
+55.26%, the other arthropods with +38.4%, and the total population of
soil arthropods with +54.04% in systemic groups of soil arthropods
extracted with Berlese funnel. However the population of mites extracted
with Berlese funnel was reduced with 80%. From results it could
concluded that glyphosate herbicide reduced all soil arthropods systemic
groups except mites collected with pitfall traps. However, it considerable
reduction effect was observed in mite populations collected with Berlese
funnel after application of glyphosate in tomato non organic system. This
reduction may be as a result of less food availability (e.g. plant roots) and
decreased green plant cover.
Out Crossing, Heterozygosis and Inbreeding with Environments Interaction in R...paperpublications3
Abstract:The progenies of five sorghum heterozygous populations’ cycles were tested under main and off-season on two different environments irrigated and rainfall conditions for their outcrossing, heterozygosity and inbreeding coefficient using SSR markers,the marker combinations were optimized according to their fragment size. Multi-locus outcrossing rate (tm) and average single-locus (ts) outcrossing rates were estimated using the MLTR software, and TFPGA computer program. The outcrossing rate effected directly by the temperature and relative humidity (RH) during the initial flowering period, which the low temperature with high RH under main season is revealed positive increased in outcrossing than off-season. Progenies outcrossing rate revealed same trend with main population outcrossing and the same trend was observed heterozygosity with decreased in inbreeding coefficient. Higher levels in outcrossing rate and heterozygosity was detected under rainfall environment in two based population progenies, but in three advanced population cycles the outcrossing rate was higher under irrigated than rainfed environment. Inbreeding coefficient revealed negative relation with outcrossing rate and heterozygosity in different population’s progenies.
Out Crossing, Heterozygosis and Inbreeding with Environments Interaction in R...paperpublications3
Abstract:The progenies of five sorghum heterozygous populations’ cycles were tested under main and off-season on two different environments irrigated and rainfall conditions for their outcrossing, heterozygosity and inbreeding coefficient using SSR markers,the marker combinations were optimized according to their fragment size. Multi-locus outcrossing rate (tm) and average single-locus (ts) outcrossing rates were estimated using the MLTR software, and TFPGA computer program. The outcrossing rate effected directly by the temperature and relative humidity (RH) during the initial flowering period, which the low temperature with high RH under main season is revealed positive increased in outcrossing than off-season. Progenies outcrossing rate revealed same trend with main population outcrossing and the same trend was observed heterozygosity with decreased in inbreeding coefficient. Higher levels in outcrossing rate and heterozygosity was detected under rainfall environment in two based population progenies, but in three advanced population cycles the outcrossing rate was higher under irrigated than rainfed environment. Inbreeding coefficient revealed negative relation with outcrossing rate and heterozygosity in different population’s progenies.
Keyword:environment, recurrent selection, outcrossing, sorghum, progenies.
Relative Abundance and ethnomedicinal Uses of some Plant Species found in Fed...AI Publications
This research was aimed to identify and determine the ethnomedicinal potential as well as the relative abundance of some selected plants in Federal University Dutsin-ma permanent site. A total of 40 plants were collected and identified from four different sites (behind senate building, Faculty of Science, Faculty of Agriculture and hostel area). In every study site, 30quadrats of 10 m X 10 m (100 sq m) size were randomly laid to study trees, herbsand shrubs species. The tree species includes all the saplings, poles and trees present in the study area. The shrubs and herbs species were studied by laying 50 quadrats of 1m X 1m (1sq m) size randomly in each study site. A totalof 33 plants were found to possess medicinal history, the plants were identified using morphological features into trees, shrubs and herbs. The total density, frequency and relative abundance of plant species collected behind senate building (Federal University Dutsin-ma) was found to be 340/ha, 260 and 26 respectively. In Faculty of Science, the total density, frequency and relative abundance of plant species collected was found to be 340/ha, 240 and 24 respectively. The total density, frequency and relative abundance of plant species collected around Faculty of Agriculture was 420/ha, 280 and 28 respectively, while at hostel area, the total density, frequency and relative abundance of plant species collected was found to be 420/ha, 280 and 28 respectively. Different plants species were collected, identified and found to possess some medicinal properties, these plants includes Sclerocaryabirrea, Sida alba, Euphorbia hirta, Sennaoccidentalis, Acacia ataxacantha, Sennaobtusifoliaand Cleome monophylla.
Wheat-pea intercropping for aphid control: from laboratory tritrophic approac...InternationalNetwork
Intercropping is an interesting practice to promote the sustainable control of insect pests such as aphids. In particular, volatile organic compounds emitted by aphid-infested intercropped plants may deter other aphid species from their host plants, while attracting natural enemies. In this study, olfactometer and net-cage behavioural assays were first conducted to determine the effect of wheat-pea mixtures combined with aphid infestations on odour preferences of the wheat aphid Sitobion avenae and two associated predator species, the ladybird Harmonia axyridis and the hoverfly Episyrphus balteatus. Healthy wheat plants were preferred by S. avenae, while wheat-pea mixtures combined with aphid infestations were significantly less attractive. H. axyridis preferred odours from healthy wheat plants mixed with aphid-infested pea plants. As for E. balteatus, their searching and oviposition behaviours were stimulated by the different wheat/pea combinations associated with aphid infestations. A field trial was also carried to compare the effect of mix and strip cropping wheat with pea on aphids and their natural enemies with both monocultures. Wheat and pea aphid populations were significantly reduced by both types of intercropping when compared to monocultures. Moreover, higher abundances of hoverflies, lacewings and ladybirds were found in wheat mixed with pea field, followed by strip cropping and monocultures. These findings show that wheat-pea intercropping can be efficient to reduce aphid populations, namely by promoting their biological control.
Benefits Over Traditional Wired Sensing Technology
Hannah's ISS
1. Independent Study Semester
How do habitat restorations affect plants and pollinators in Metro Vancouver Parks?
Hannah Gehrels
For Elizabeth Elle and David Green
December 9, 2013
Abstract
Anthropogenic disturbance associated with urban growth facilitates the spread of invasive
plant species which compete with native species for access to mutualists such as pollinators.
Habitat restorations are often purposed to prevent native species loss and restore mutualistic
interactions. We used 12 semi-urban parks (each with a restored and control plot) located
throughout the Greater Vancouver area to investigate how habitat restorations via native
plantings and invasive plant removals impact plant and pollinator communities. We found that
plant richness and Simpson’s diversity were higher in restored plots, but that abundance was
similar with restoration treatment. We also found that pollinator richness was higher in restored
plots when we controlled for time, and that abundance and Simpson’s diversity tended to follow
this same trend. Finally, we found that nestedness and asymmetry were higher in control
(invaded) plots due to a higher abundance of generalist invasive plants, suggesting that invasive
plants become highly integrated into plant-pollinator networks and may increase network
resilience. We suggest that when invasive plants are removed, managers should ensure that
alternative forage is available by planting generalist native species that have radially symmetrical
flowers and that together provide floral resources across the appropriate phenological and spatial
scales.
Key words: invasive species, restorations, floral resources, plant-pollinator networks
2. Introduction
Anthropogenic disturbance associated with urban growth facilitates the spread of invasive
species which modify the structure and stability of communities (Hierro et al. 2006, Kneitel and
Perrault 2006). In particular, invasive plants alter the composition of native plant communities
by outcompeting native species for nutrients, water and light (Abraham et al. 2009). Invasive
plant species can also compete with native species for access to mutualists such as pollinators or
seed dispersers (Traveset and Richardson 2006), and generally will affect animals that depend
on plants for food and habitat (Litt and Steidl 2010).
Ecological restorations are increasingly used in attempts to prevent native species loss
and reestablish ecosystem function. Restorations return the species composition and physical
structure of disturbed habitat to a goal state (often based on historic conditions; SER Working
Group 2004). Restorations of anthropogenically altered terrestrial habitats often involve planting
native species, sometimes coupled with removal of invasive species. Increased species richness
or abundance is frequently used as a measure of restoration success, but species richness does not
always capture ecosystem function (Elle et al. 2012). Pollinators play an important role in
providing ecosystem services and may serve as a good indicator of ecosystem recovery
(Montoya et al. 2012), but a consideration of plant-pollinator interactions as well as changes in
richness or abundance is required (Elle et al. 2012).
Plant-pollinator interaction networks are useful because they focus on the functional role
of species in a community rather than simple presence/absence (Elle et al. 2012). Since
pollinators are essential for the reproduction of the vast majority of flowering plants (Ollerton et
al. 2011), evaluating plant-pollinator interactions can provide information about the resilience of
communities. Plant-pollinator networks have a nested structure whereby interactions are
organized around a core group of generalist plant and pollinator species (Kaiser-Bunbury et al.
2011). Some specialized pollinators visit generalist plants and vice versa resulting in
asymmetrical dependence (Elle et al. 2012). More nested, asymmetrical and generalized
interaction networks tend to be more resilient in the face of further disturbance (Thébault and
Fontaine 2010, Elle et al. 2012).
Mixed results have been found in studies that investigated impacts of restoration on
pollinators in urban and semi-urban environments. Some studies have shown that habitat
restorations can increase pollinator abundance, richness and diversity (Carvell et al. 2007,
Hopwood 2008), whereas others have found no effect (Forup and Memmott 2005, Bartomeus et
al. 2008, Matteson and Langellotto 2011, Williams 2011, Ferrero et al. 2013). Most studies agree
that invasive plants can be can become high integrated into plant-pollinator interaction networks
(Memmott and Waser 2002, Morales and Aizen 2006, Lopezaraiza-Mikel et al. 2007,
Valdovinos et al. 2009, McKinney and Goodell 2010), which serve to increase nestedness and
asymmetry (Aizen et al. 2008, Bartomeus et al. 2008). Given this variation in results, the utility
of restoration efforts should be considered in different ecosystem types. In addition, none of
these studies explicitly investigated how pollinators change over the season. The phenology of
flowering plants (available forage) in a community is important because many solitary bee
species are only actively foraging for a few weeks (O’Toole and Raw 2004, Godoy et al. 2009).
The timing of native plant availability may be crucial when restoring plant communities to
support pollinators, and when assessing restoration success.
Here we aim to investigate the effects of native plant restorations coupled with the
removal of invasive species on the abundance, richness and diversity of pollinators and the
resilience of plant-pollinator interactions in urban parks in the Greater Vancouver Regional
3. District. We hypothesize that in comparison to the control areas, the restored areas will have (1)
higher pollinator abundance, richness and diversity, (2) higher floral abundance, richness and
diversity, (3) more temporally uniform flower and pollinator abundance and richness, and (4)
lower nestedness and asymmetry, and a higher specialization index.
Methods
Study Area
We conducted this study in 12 urban to semi-urban parks throughout the Greater
Vancouver area from April 24-August 22, 2013 (Fig. 1). Each site included a pair of plots, one
restored and one control with plots matched in shape and area. Plots were an average of 176m
apart from each other (range: 50-440m, Table 1). We also sampled at a 13th site (Lower Seymour
Conservation Reserve), and a third plot at CF-2, but did not include these plots in our analysis
because the paired plots were not similar enough for an accurate comparison between restored
and control plots.
Vegetation sampling
Only potentially pollinator-attractive plants with open flowers were sampled for this
research, and so grasses, ferns, etc were not included. Flower abundance, richness and diversity
for each plot were sampled on the same days pollinators were assessed. For the seven sites that
had plots with linear hedgerows, vegetation was sampled along a 50 m transect along the
hedgerow edge, with samples taken at 1-m intervals. The line intercept method was used such
that the number of open flowers intersecting a 1m line perpendicular to the transect (into the
hedgerow) was counted by species. For the five sites that had plots with approximately
rectangular areas, the vegetation was sampled using the same line intercept method, but at
regular intervals along 5 parallel transects placed in a stratified random manner. Length of
transect varied with the size of the plot. Densely clustered floral heads (e.g. in families
Asteraceae, Brassicaea, Plantaginaceae) were considered a single “flower” for the purposes of
this study (see Appendix A for floral unit designations by species).
Pollinator sampling
We caught floral visitors (hereafter pollinators) with hand-nets directly from flowers. We
sampled each site approximately every two weeks on warm, sunny days (temperature ≥ 14, low
wind, and sunny to partly cloudy). Pollinators were collected for 15 minutes by each of two
people (= 30 minutes per plot per sample date), in the morning (1000 – 1200h), midday (1200 –
1400h) or late afternoon (1400 – 1600h). Paired plots were sampled on the same day, and most
sites were sampled three times in each of the three times of day, for a total of 9 sample episodes
(4.5 hours) per site. Three of the sites (BB-1, BB-2, and OM) were not restored or accessible
until after we had started sampling, so were sampled for 7 sample episodes only (3.5 hours).
Flower species identity was noted for each pollinator collected. All bees were identified to
species except those for which revised keys were not available. Flies and wasps were identified
at least to family, but to genus or species where possible.
4. Statistical analysis
We compared abundance, species richness, and evenness of plants and pollinators in
restored and control plots. To examine species evenness, we calculated Simpson’s diversity
index for each plot within a site: 𝐷 = 1 − (∑
𝑛𝑖
𝑁
)
2
, where ni is the number of individuals of species
i, and N is the total number of individuals. To investigate differences in plant and pollinator
abundance, richness, and Simpson’s diversity between restored and control plots, we performed
a mixed effects ANOVA with treatment as the main effect and site as a random effect. To
compare pollinator and floral abundance and richness over time between the restored and control
plots, we performed a repeated measures ANOVA with time, treatment, and their interaction in
the model, again including site as a random effect. We could not compare Simpson’s diversity
over time because on some date/site combinations, no pollinators were caught, resulting in
undefined values. To examine the functional shifts in plant and pollinator communities and to
identify interactions that may be vulnerable to disturbance, we created a plant-pollinator network
for each plot. Using the bipartite package in R (Dormann et al. 2008), we calculated nestedness,
asymmetry, and the specialization index (H2’; Blüthgen et al. 2006), for each network and
compared between restored and control plots using a mixed effects ANOVA with site as a
random variable.
Results
Plant abundance, richness and diversity
Plant abundance did not differ with restoration treatment (F1,11=0.06, P =0.81, Fig. 2).
Richness and Simpson’s diversity were significantly higher in restored plots compared to control
plots (richness: F1,11=5.62, P =0.04, Simpson’s diversity: F1,11=5.97, P =0.03). Repeated
measures analysis indicated that abundance and richness changed over time with abundance
peaking in early July and richness peaking in late July (abundance: F8,174=5.37, P<0.0001,
richness: F8,174=4.78, P<0.0001), although the interaction term was once again not significant
(abundance: F8,173=0.70, P=0.69, richness: F8,171=1.21, P=0.30). Richness was significantly
higher in the restored plots in this analysis, whereas the difference in abundance remained non-
significant (richness: F1,173=17.30, P<0.001, abundance F1,173=0.09, P=0.77). However, the
restored plots tended to have higher flower abundance early and late in the season, whereas the
control plots tended to have higher abundance in the middle of the season driven by a high
abundance of invasive Rubus discolor in several of the control plots (Fig. 3).
Pollinator abundance, richness and diversity
We netted 3247 individuals in total, representing 150 species (bees: 24 Halictidae, 21
Megachilidae, 17 Andrenidae, 10 Bombus, 10 other bees, as well as the highly managed Apis
mellifera; flies: 41 Syrphidae, 9 other flies; 17 wasps; and 8 other floral visitors (hummingbird,
butterflies, beetle, etc)). Although all tended to be higher in the restored plots (Fig. 2), pollinator
abundance, richness, and Simpson’s diversity did not significantly differ with restoration
treatment (abundance: F1,11=1.74, P=0.21, richness: F1,11=1.53, P=0.24, and Simpson’s
diversity: F1,11=0.88, P=0.37). The repeated measures analysis indicated that abundance and
richness changed over time (abundance: F8,173=2.10, P=0.04, richness: F8,173=3.41, P=0.001).
Abundance in the restored plots peaked in the third sampling period (late May), which seems to
be driven by 2 species: the solitary Andrena miserabilis which was only found at one plot (PS-2)
5. on one day, and the highly managed Apis melifera at another plot (AG) where managed hives
were kept nearby. Abundance in the control plots and richness for both treatments peaked in late
July (Fig. 4). There was no difference in how control vs. restored plots responded to time
(interaction term was not significant; abundance: F8,171=0.34, P=0.95, richness: F8,171=0.61,
P=0.77). Both abundance and richness were higher in the restored plots in this analysis, but only
richness achieved significance (abundance: F1,171=2.92, P=0.089, richness: F1,171=7.10,
P=0.008).
Network metrics
Nestedness tended to be higher in the control plots, but did not achieve statistical
significance (F1,11=4.05, P =0.07, Fig. 5). Asymmetry was significantly higher in the control
plots compared to the restored plots (F1,11=11.45, P =0.01). The specialization index (H2’) did
not differ with restoration treatment (F1,11=0.39, P =0.54).
Discussion
Plants and Pollinators
Restorations increased floral species richness and Simpson’s diversity as we predicted,
but these improved floral resources did not translate into higher abundance, richness or diversity
of pollinators in the restored plots. In general, pollinator communities are expected to track plant
communities that provide food resources (Potts et al. 2004, Hennig and Ghazoul 2011). Some
previous studies found that restoration increased pollinator richness (Carvell et al. 2007,
Hopwood 2008). Of those studies that did not find increased richness with restoration (Forup and
Memmott 2005, Bartomeus et al. 2008, Matteson and Langellotto 2011, Williams 2011, Ferrero
et al. 2013), there were three main interpretations that apply to our study which we discuss
below.
First, native plant additions may have been insufficient numerically to produce a
measureable increase in pollinator species richness (Matteson and Langellotto 2011). In our
study, restorations improved floral richness and Simpson’s diversity, but floral abundance was
similar with restoration treatment. The most common invasive plant species, Himalayan
Blackberry (Rubus discolor), produces large numbers of flowers per unit area. It seems that the
native plant additions in the restored plots were insufficient in number to produce a measurable
increase in floral abundance relative to the floral resources in the control plots from abundant
invasive species like Rubus discolor. This similarity in floral abundance with restoration
treatment may have, in turn, affected our lack of significant improvement in pollinator species
richness. A model by Matteson and Langellotto (2011) suggested that it would take
approximately 200-250 flowers to increase bee richness by one species. If the assumptions of
this model apply to our system, the similarity in flower abundance with restoration treatment that
we found in our experiment would be predicted to be associated with no difference in pollinator
richness, as we found.
Second, pollinator community composition may be different even though abundance,
richness and diversity are similar (Williams 2011, Ferrero et al. 2013, Wray et al. in press). In
our study, pollinators could have flown between plots (plots were on average 176m apart
whereas the typical flight distance of solitary bees is 200-400m, Greenleaf et al. 2007), which
may have dampened the observed differences in pollinator communities between restored and
6. control plots. However, the ‘floral market’ hypothesis suggests that pollinators choose between
plant species on the basis of the quality of their resources (nectar and pollen, Chittka and
Schürkens 2001). This hypothesis indicates that even if pollinators were flying between plots,
they were making choices about which flowers to visit, so we could still expect a difference in
pollinator communities if the plant communities are different. For instance, we might expect
more generalist pollinator species to be present in the control plots that have more generalist
invasive plants (Cane et al. 2006). In future analyses of our data, we will pursue community-
scale analyses (e.g. ordination) to assess differences in the plant and pollinator communities,
rather than just evaluating differences in richness and diversity (Wray et al. in press).
Thirdly, flowering plants do not provide all the resources needed for pollinators. Nesting
sites for soil-nesting bees may be particularly limited in urbanized landscapes due to soil
compaction and pavement (Cane et al. 2006, Matteson et al. 2008). In contrast to floral
resources, however, nesting resource availability is difficult to assess, and further studies on bee
nest site use in urban areas are required. If nest sites are a major limiting factor for some
pollinators, simple additions of floral resources may not be enough to increase overall pollinator
abundance, richness and diversity (Potts et al. 2005).
Plant and pollinator abundance and richness varied with time, peaking in mid-season.
The variation over time did not vary with restoration treatment, however, indicating that floral
resource availability (and the abundance and richness of pollinators requiring those resources)
was similar in control and restoration plots. This is important because pollinators rely on the
overlap between their flight periods and the flowering periods of each plant species in a
particular area (Bosch et al. 1997, Basilio et al. 2006). One interesting finding was that when
seasonality was controlled in the repeated measures analysis, pollinator richness was
significantly higher in restored plots. That is, controlling for the variability among sample
periods allowed us to detect that pollinator richness was improved with restoration. The increase
in richness is most pronounced in the late season, which may have been caused by the similar
increase in plant richness at this time. It may be useful to consider floral availability at other
times of the season and whether restoration planning could be improved by considering plant
flowering time.
Plant-pollinator networks
Asymmetry and nestedness were higher (asymmetry significantly so) in control plots.
This finding suggests that control (invaded) plots are more resilient to disturbance than restored
plots. Many studies agree that invasive plant species can become highly integrated into plant-
pollinator interaction networks (Memmott and Waser 2002, Morales and Aizen 2006,
Lopezaraiza-Mikel et al. 2007, Valdovinos et al. 2009, McKinney and Goodell 2010). Aizen et
al. (2008) found that more invaded sites had higher asymmetry than their less invaded
counterparts resulting from a transfer in the plant-pollinator interactions from generalist native
plant species to super-generalist invasive plant species. These generalist invasive plants reduce
the average interaction strength in the network and increase nestedness and asymmetry (Aizen et
al. 2008, Bartomeus et al. 2008, Valdovinos et al. 2009). Since most of the nonnative plants in
our study area were generalist plants with radially symmetrical flowers that allow any insect to
interact with them (e.g. Rubus discolor, Hypochaeris radicata, and Ranunculus repens), this
reason seems to fit for our study as well. In contrast, several native plant species used in the
restorations had limited pollinator access (e.g. Lonicera involucrata, Ribes sanguineum and
Lupinus arcticus), and as such, were not available to all pollinators.
7. High asymmetry and nestedness are generally thought to confer higher network stability
in the face of disturbance due to more redundant plant-pollinator interactions (Elle et al. 2012).
However, a focus on these metrics may overlook other subtle changes in network structure. For
example, Aizen et al. (2008) found that invasive plant species decreased the amount of native-
native interactions, some of which may be ecologically and evolutionarily important. For this
reason, it is important to investigate how plant-pollinator interactions change with habitat
restorations, in addition to calculating these network parameters.
Since native plants coevolved with native pollinators, specialized interactions may have
formed over evolutionary time, increasing the amount of specializations in a network composed
of primarily native species compared to an area with invasive plants (Gotlieb et al. 2011). In our
study, however, the specialization index (H2’) did not differ with restoration treatment even
though restored sites had fewer invasive species than control sites (20.25% of the flowers in
restored plots were invasive compared to 47.92% in control plots). It is possible that the
differences in the amount of invasive species between plots may not have been large enough to
have a measureable impact on the change in specialization index. Additionally, our analyses
were created using cumulative networks which groups all of the plant and pollinators together,
including species that are not active at the same time(Basilio et al. 2006). This method
exaggerates generalization scores and could overlook possible changes in the degree of
specialization over the season (Basilio et al. 2006, Burkle and Alarcón 2011). We suggest that
further analyses of plant-pollinator networks include intra-annual variation.
Conclusions
Invasive plants are considered to be the third major cause of pollinator diversity loss
(Kearns et al. 1998). Our study suggests that invasive plants do have a negative impact, but that
the effect may not be as negative as previously thought. Specifically, our data show that
pollinator richness increased with restorations via native plantings when controlling for time, and
that pollinator abundance and Simpson’s diversity tended to be higher in the restored plots as
well. However, the integration of invasive plant species into native networks may actually serve
to make the native network more robust and resilient to changes in species composition
(Memmott et al. 2004, Ferrero et al. 2013).
Our study shows that invasive plants become highly integrated into plant-pollinator
networks, which has implications for managers. Specifically, we suggest that when removing
invasive flowering plants, care should be taken to ensure that alternative forage is available for
the pollinators that rely on those invasive plants within the appropriate phenological and spatial
scales. Habitat restorations that involve planting native species should incorporate combinations
of flowering plants that together provide a continuous source of floral resources for pollinators
over the course of the season. Additionally, we suggest that flower morphology should also be
considered in habitat restoration plans. Generalist native plants that have flowers with radial
symmetry (e.g. Symphoricarpos albus, Rosa nutkana, and Rubus spectabilis in our study area)
may serve to increase pollinator richness and overall network resilience.
Acknowledgements
Angela Fortune assisted with field and lab work, and Jennifer Avery assisted with the
plant analysis. Elizabeth Elle and David Green provided comments and supervision. Funding
was provided by Metro Vancouver, the Environmental Youth Alliance, Simon Fraser University
Biology department, and the Natural Sciences and Engineering Council (NSERC) of Canada.
8. Works Cited
Abraham, J. K., J. D. Corbin, and C. M. D. Antonio. 2009. California native and exotic perennial
grasses differ in their response to soil nitrogen, exotic annual grass density, and order of
emergence. Plant Ecology 201:445–456.
Aizen, M. A., C. L. Morales, and J. M. Morales. 2008. Invasive mutualists erode native
pollination webs. Plos Biology 6:396–403.
Bartomeus, I., M. Vilà, and L. Santamaría. 2008. Contrasting effects of invasive plants in plant-
pollinator networks. Oecologia 155:761–70.
Basilio, A. M., D. Medan, J. P. Torretta, and N. J. Bartoloni. 2006. A year-long plant-pollinator
network. Austral Ecology 31:975–983.
Bosch, J., J. Retana, X. Cerdá, and S. Url. 1997. International Association for Ecology Flowering
Phenology, Floral Traits and Pollinator Composition in a Herbaceous Mediterranean Plant
Community. Pecologia 109:583–591.
Burkle, L. a, and R. Alarcón. 2011. The future of plant-pollinator diversity: understanding
interaction networks across time, space, and global change. American Journal of Botany
98:528–38.
Cane, J. H., R. L. Minckley, L. J. Kervin, T. H. Roulston, and N. M. Williams. 2006. Complex
responses within a desert bee guild (Hymenoptera: Apiformes) to urban habitat
fragmentation. Ecological applications : a publication of the Ecological Society of America
16:632–44.
Carvell, C., W. R. Meek, R. F. Pywell, D. Goulson, and M. Nowakowski. 2007. Comparing the
efficacy of agri-environment schemes to enhance bumble bee abundance and diversity on
arable field margins. Journal of Applied Ecology 44:29–40.
Chittka, L., and S. Schürkens. 2001. Successful invasion of a floral market. Nature 411:653.
Dormann, C. F., G. B, and F. J. 2008. Introducing the bipartite Package: Analysing Ecological
Networks. R news 8:8–11.
Elle, E., S. L. Elwell, and G. A. Gielens. 2012. The use of pollination networks in conservation.
Botany 90:525–534.
Ferrero, V., S. Castro, J. Costa, P. Acuña, L. Navarro, and J. Loureiro. 2013. Effect of invader
removal: pollinators stay but some native plants miss their new friend. Biological Invasions
15:2347–2358.
Forup, M. L., and J. Memmott. 2005. The Restoration of Plant-Pollinator Interactions in Hay
Meadows. Restoration Ecology 13:265–274.
Godoy, O., D. M. Richardson, F. Valladares, and P. Castro-Díez. 2009. Flowering phenology of
invasive alien plant species compared with native species in three Mediterranean-type
ecosystems. Annals of Botany 103:485–94.
Gotlieb, A., Y. Hollender, and Y. Mandelik. 2011. Gardening in the desert changes bee
communities and pollination network characteristics. Basic and Applied Ecology 12:310–
320.
Greenleaf, S. S., N. M. Williams, R. Winfree, and C. Kremen. 2007. Bee foraging ranges and
their relationship to body size. Oecologia 153:589–96.
Hennig, E. I., and J. Ghazoul. 2011. Pollinating animals in the urban environment. Urban
Ecosystems 15:149–166.
9. Hierro, J. L., D. Villarreal, Ö. Eren, J. M. Graham, and R. M. Callaway. 2006. Disturbance
Facilitates Invasion: The effects are stronger abroad than at home. The American Naturalist
168:144–156.
Hopwood, J. L. 2008. The contribution of roadside grassland restorations to native bee
conservation. Biological Conservation 141:2632–2640.
Kaiser-Bunbury, C. N., T. Valentin, J. Mougal, D. Matatiken, and J. Ghazoul. 2011. The
tolerance of island plant-pollinator networks to alien plants. Journal of Ecology 99:202–
213.
Kneitel, J. M., and D. Perrault. 2006. Disturbance-induced changes in community composition
increase species invasion success. Community Ecology 7:245–252.
Litt, A. R., and R. J. Steidl. 2010. Insect assemblages change along a gradient of invasion by a
nonnative grass. Biological Invasions 12:3449–3463.
Lopezaraiza-Mikel, M. E., R. B. Hayes, M. R. Whalley, and J. Memmott. 2007. The impact of an
alien plant on a native plant-pollinator network: an experimental approach. Ecology letters
10:539–50.
Matteson, K. C., J. S. Ascher, and G. A. Langellotto. 2008. Bee Richness and Abundance in New
York City Urban Gardens Bee Richness and Abundance in New York City Urban Gardens.
Conservation Biology and Biodiversity 101:140–150.
Matteson, K. C., and G. A. Langellotto. 2011. Small scale additions of native plants fail to
increase beneficial insect richness in urban gardens. Insect Conservation and Diversity
4:89–98.
McKinney, A. M., and K. Goodell. 2010. Plant–pollinator interactions between an invasive and
native plant vary between sites with different flowering phenology. Plant Ecology
212:1025–1035.
Memmott, J., and N. M. Waser. 2002. Integration of alien plants into a native flower-pollinator
visitation web. Proceedings. Biological sciences / The Royal Society 269:2395–9.
Montoya, D., L. Rogers, and J. Memmott. 2012. Emerging perspectives in the restoration of
biodiversity-based ecosystem services. Trends in Ecology and Evolution 27:666–672.
Morales, C. L., and M. a. Aizen. 2006. Invasive mutualisms and the structure of plant-pollinator
interactions in the temperate forests of north-west Patagonia, Argentina. Journal of Ecology
94:171–180.
O’Toole, C., and A. Raw. 2004. Bees of the world. Pages 1–192. Cassell Illustrated, London.
Ollerton, J., R. Winfree, and S. Tarrant. 2011. How many flowering plants are pollinated by
animals? Oikos 120:321–326.
Potts, S. G., B. Vulliamy, S. Roberts, C. O’Toole, a Dafni, G. Ne’Eman, and P. Willmer. 2005.
Role of nesting resources in organising diverse bee communities in a Mediterranean
landscape. Ecological Entomology 30:78–85.
Potts, S. G., B. Vulliamy, S. Roberts, C. O’Toole, A. Dafni, G. Ne’eman, and P. G. Willmer.
2004. Nectar resource diversity organises flower-visitor community structure. Entomologia
Experimentalis et Applicata 113:103–107.
Thébault, E., and C. Fontaine. 2010. Stability of ecological communities and the architecture of
mutualistic and trophic networks. Science 329:853–6.
Traveset, A., and D. M. Richardson. 2006. Biological invasions as disruptors of plant
reproductive mutualisms. Trends in ecology & evolution 21:208–16.
Valdovinos, F. S., R. Ramos-Jiliberto, J. D. Flores, C. Espinoza, and G. López. 2009. Structure
and dynamics of pollination networks: the role of alien plants. Oikos 118:1190–1200.
10. Williams, N. M. 2011. Restoration of Nontarget Species: Bee Communities and Pollination
Function in Riparian Forests. Restoration Ecology 19:450–459.
11. Figures and Tables
Figure captions:
Figure 1: Map of study sites. Each site included a pair of plots, one restored and one non-
restored.
Figure 2: Plant and pollinator abundance, richness, and Simpson’s diversity index for restored
and non-restored plots averaged across sites. * indicates marginally non-significant results
(p<0.10), and ** indicates significant results (p<0.05).
Figure 3: Average floral abundance and richness over time.
Figure 4: Average pollinator abundance and richness over time
Figure 5: Nestedness, asymmetry, and the Specialization index (H2) for restored and non-
restored plots averaged across sites. * indicates marginally non-significant results (p<0.10), and
** indicates significant results (p<0.05).
14. 0
1
2
3
4
5
6
7
8
9
10
Late AprilEarly May Late May Early
June
Late June Early July Late July Early
August
Late
August
Averagefloralspeciesrichnesspersite
Time
0
50
100
150
200
250
300
350
Averagenumberofflowerspersite
Restored
Control
17. Table 1: List of Sites
Site Site Code Plot Shape UTM
Year of
restoration
Distance
between
plots
Delta Heritage Air
Park
DP
Hedgerow
49°04’44.15”N
122°56’16.16”W
2005 120m
Centennial Beach BB-1 Polygon
49°00’57.15”N
123°02’28.33”W
2013 165m
Boundary Bay
Regional Park
BB-2 Polygon
49°01’02.55”N
123°03’07.59”W
Late 1990s 380m
Pacific Spirit Park-
Camosun Bog
PSC Polygon
49°15’13.54”N
123°11’46.24”W
2010 30m
Pacific Spirit Park
(near the Museum of
Anthropology)
PSM Hedgerow
49°16’14.86”N
123°15’32.20”W
2006-2007 440m
Oak Meadows Park OM Polygon
49°14’17.67”N
123°07’34.34”W
2013 70m
Aldergrove Regional
Park
AG
Hedgerow
49°00’34.21”N
122°27’03.72”W
2002-2003 100m
Brae Island Regional
Park
BI
Hedgerow
49°10’30.90”N
122°34’48.55”W
2007 190m
Campbell Valley
Regional Park
CV
Hedgerow
49°01’03.15”N
122°39’48.44”W
2002-2004 60m
Colony Farm
Regional Park-2
CF-2
Hedgerow
49°14’21.46”N
122°47’49.64”W
2007 300m
Colony Farm
Regional Park-1
CF-1
Hedgerow
49°14’27.41”N
122°48’30.74”W
1999 120m
Tynehead Regional
Park
TH
Polygon
49°11’02.30”N
122°44’59.36”W
2012 260m