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Juz 024
Juz 024
doi: 10.1093/jue/juz024
Research article
Submitted: 18 June 2019; Received (in revised form): 31 October 2019; Accepted: 11 December 2019
Abstract
Green roofs are increasingly promoted for urban biodiversity conservation, but the value of these novel habitats is uncer-
tain. We aimed to test two hypotheses: (i) green roofs can support comparable invertebrate family and order richness, com-
position and abundances to ground-level habitats and (ii) green roofs planted with native species from local habitats will
support a richer invertebrate community at family and order level than other green roofs. We sampled the invertebrate
community on green roofs dominated by native grassland or introduced succulent species in Melbourne, Australia, and
compared these to the invertebrate community in ground-level sites close by, and sites with similar vegetation types. The
only significant differences between the invertebrate communities sampled on green roofs and ground-level habitats were
total abundance and fly family richness, which were higher in ground-level habitats. Second hypothesis was not supported
as invertebrate communities on green roofs supporting a local vegetation community and those planted with introduced
Sedum and other succulents were not detectably different at family level. The per cent cover of green space surrounding
each site was consistently important in predicting the richness and abundance of the invertebrate families we focussed on,
while roof height, site age and size were influential for some taxa. Our results suggest that invertebrate communities of
green roofs in Melbourne are driven largely by their surrounding environment and consequently the effectiveness of green
roofs as invertebrate habitat is highly dependent on location and their horizontal and vertical connection to other habitats.
Key words: green infrastructure, pollinators, urban bees, arthropods, urban greening
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2 | Journal of Urban Ecology, 2020, Vol. 6, No. 1
suitable habitat inhospitable. The relative isolation of a green found green roofs to be more biodiverse than conventional roofs
roof may allow for plant species and animals sensitive to distur- (Pearce and Walters 2012; Partridge and Clark 2018; Belcher
bance, or prone to competitive exclusion, to persist. Green roofs et al. 2019). Somewhat surprisingly, relatively few studies have
have therefore been proposed as components of urban bio- undertaken more conservation relevant comparison of inverte-
sphere networks (Kim 2004), as sites that could support metapo- brate communities on extensive green roofs to those found in
pulations of endangered butterflies in urban areas (Snep, nearby equivalent ground-level habitats that have similar vege-
WallisDeVries, and Opdam 2011) and replacement habitats for tation (for exceptions, see Ksiazek, Fant, and Skogen 2012;
biodiverse areas destroyed by development (Lorimer 2008). Braaker et al. 2014, 2017). There has also been little evaluation
They are also included in biodiversity policy recommendations of green roofs specifically designed for biodiversity (Butler,
The selected green roofs were either planted with a range of pan traps, which are widely used for sampling agricultural pests
native forb and grass species from the critically endangered (Southwood 1978), and to sample the abundance and diversity
grasslands to the north and west of Melbourne (sites 1, 2 and 3), of a variety of insect communities (Leong and Thorp 1999) pre-
which are considered a habitat template for south-eastern dominantly sample flying invertebrates. This combination of
Australian green roofs (Williams, Rayner, and Raynor 2010; methods allowed us to survey diurnal and nocturnal inverte-
Rayner et al. 2016), or with succulents, primarily introduced brates (Hill 2005). The sampling design aimed to capture a com-
Sedum species (sites 4, 5 and 6) (Supplementary Table S1). All parative sample between sites without assessing the full
green roofs sampled had a scoria (lava rock)-based growing me- assemblage composition within sites.
dium, similar to that described by Farrell et al. (2012), that was Three 30-ml tube pitfall traps were installed in a line 50 cm
was considered a random effect (to indicate which of the six had a significantly greater richness of Diptera (flies) than green
site clusters each sample was associated with) and ‘type’ a fixed roofs (P ¼ 0.02), however companion and green roof sites had
effect. Type included green roofs, companion and nearby site comparable Diptera richness (P > 0.05). No other significant dif-
types. All response variables were continuous. Order abundance ferences were found between site types for Coleoptera (beetles),
and beetle family richness data were log transformed to im- Hymenoptera (bees, wasps, ants) or Hemiptera (true bugs) fam-
prove the normality of the data. Models were run using a ily richness (P > 0.05 in all cases). There was a marginally greater
Gaussian distribution for total abundance (log), Coleoptera, Coleoptera family richness on grassland roofs than succulent
Diptera, Hemiptera and Hymenoptera family richness, pollina- roofs (P ¼ 0.047), where only one of the three succulent roofs
tor abundance (log) and a Poisson distribution for ordinal rich- were found to contain beetles. No differences between family
Taxon Median (Min, max) Median (Min, max) Median (Min, max) Total
Collembola 31.5 (0, 690) 1580.5 (5, 11 496) 1134.5 (2, 2553) 25 457
Diptera 49.5 (19, 87) 90.5 (20, 153) 258 (69, 772) 2734
Hymenoptera 63 (7, 153) 89 (62, 498) 81.5 (21, 191) 1961
Hemiptera 37 (4, 185) 38.5 (5, 824) 25.5 (13, 45) 1485
Coleoptera richness 1 0 2 3 0 5 3 0 9 12
Hymenoptera richness 3.5 2 5 5 1 5 4 1 6 19
Hemiptera richness 3.5 2 4 3 1 5 2 1 4 6
Diptera richness 11 10 16 15 7 21 18 12 24 35
invertebrates and she notes this was probably due to a high and richness of native bees found in central Melbourne (Mata
number of snails introduced on the pre-grown sedum mats. et al. 2015) where most of the green roofs sampled were located.
We found no difference in the order-level composition of Few studies have compared the biodiversity of green roofs
green roof invertebrate communities when compared to adja- with different types of vegetation with the exceptions being
cent ground-level vegetation and similar ground-level vegeta- Madre et al. (2013), who investigated arthropods and Joimel
tion nearby. Of the four orders identified to family level, only et al. (2018), who sampled Collembola. Apart from marginally
the composition of fly families differed significantly between greater beetle family richness, the invertebrate communities on
green roofs and both nearby and companion ground-level habi- the native grassland green roofs, that theoretically should fa-
tats. This may be driven by a greater abundance of individuals vour Melbourne’s endemic invertebrate biodiversity, were not
in certain Diptera families at ground level, but three of four substantially different from the succulent green roofs domi-
green roofs also had a greater proportion of individuals in the nated by introduced sedum species and other exotic succulents.
family Chironomidae than the ground-level sites. Most species We expected to see a greater influence of green roof vegetation
in this family have aquatic larvae and short-lived adults that type across all invertebrate groups sampled because increased
eat nectar and pollen. Their relatively high abundances on roofs vegetation complexity and high proportion of native plant spe-
may be due to colonisation of, and probable breeding in, stand- cies is associated with higher biodiversity at ground level (Tews
ing water stored in the green roof drainage layers by individuals et al. 2004; Threlfall et al. 2017) and on green roofs (Madre et al.
originating from nearby ground-level water bodies, which in- 2013). The relatively young age of all the green roofs sampled
cluded ornamental ponds at two sites and ephemeral wetlands could mean that differences are yet to develop and long-term
at a neighbouring brownfield development site. This highlights monitoring may produce different results. Alternatively, be-
the influence of the surrounding landscape on the composition cause there are very few areas of native grassland remaining in
of green roof invertebrate communities. Melbourne, and most of these are on the city’s outskirts distant
Low numbers of native bees and hoverflies were found to from the green roofs (Williams, McDonnell, and Seager 2005),
use both succulent and grassland green roofs. However, no sig- specialist native grassland invertebrates may be unavailable to
nificant difference in pollinator richness, abundance or compo- colonise the green roofs or the small patches of grassland we
sition between green roof and ground-level sites and succulent sampled. In addition, the ground-storey vegetation in most ur-
and grassland sites was detected. This result differs from other ban parks is dominated by exotic grasses and herbs meaning
studies, such as Ksiazek, Fant, and Skogen (2012) who found that native invertebrates, particularly specialist species that uti-
that green roofs supported a lower abundance of bees than lise native ground-storey plants, may be absent from large parts
nearby urban green space but may reflect the low abundance of the urban landscape. Confirming this is difficult due to the
6 | Journal of Urban Ecology, 2020, Vol. 6, No. 1
Table 3: Strongest competing univariate linear mixed models of invertebrate communities on green roof, companion sites and nearby sitesa
a
Only models within two AICc points of the top model are listed. The sign within parentheses indicate the direction of the relationship for each model parameter.
lack of research on the distribution and composition of (Williams, Rayner, and Raynor 2010) and the oldest green roof in
Melbourne’s invertebrate communities and the effects of urban- this study was only 7 years old, and the others were between 1
isation on them, which are hampered by a lack of taxonomic ex- and 3 years. Consequently, age effects may become more appar-
pertise (Yen 2011). ent in the future. The abundance and richness of invertebrates
We found that roof height negatively influenced ordinal collected from green roofs was expected to increase with green
abundance and fly family richness. This result is similar to roof area, but area was not found to consistently influence in-
other studies, such as MacIvor (2016), who found a decline in vertebrate communities, only affecting beetle family richness.
the number of solitary bees and wasps utilising artificial nests Similar to the roof age, the range of roof areas was quite narrow
with increasing green roof height, and Madre et al. (2013), who due to the still limited number of extensive green roofs in
found height negatively affected the richness and abundance of Melbourne. Braaker et al. (2014) also found that green roof area
spiders and taxonomic composition of true bugs and beetles on was not an important determinant of their invertebrate com-
French green roofs. As green roof height increases, inverte- munities but we acknowledge that different invertebrate taxa
brates, particularly those with limited dispersal ability, may may respond to area differently.
find them increasingly difficult to colonise, leading to less di- This study sampled invertebrates on all available extensive
verse invertebrate communities compared to ground level. Our green roofs in Melbourne in spring and autumn, using standard
results also suggest that the insects that are able to disperse to methods (pitfall and pan traps) over adequate time periods [7
green roofs do not increase in abundance to numbers similar to days (once per season) and 24 h (three times each season), re-
ground-level sites. This could be because the sampled green spectively]. We recognise that replication is low due to the rela-
roofs could not provide sufficient resources for a large abun- tively few extensive green roofs in Melbourne and that using
dance of individuals due to low primary productivity caused by additional sampling methods over longer time periods would
the low nutrient inorganic substrates used. Similar processes probably have collected a greater abundance and richness of
have been shown to influence invertebrate communities in taxa. There is also potential sampling bias in our results as pan
ground-level urban habitats (Shochat et al. 2004; Threlfall, Law, traps may not effectively sample larger bees (Roulston, Smith,
and Banks 2012). and Brewster 2007) although it should be noted there are no
The per cent cover of surrounding green space was the most Bombus species on mainland Australia (Buttermore 1997).
frequent habitat variable affecting green roof invertebrate com- Notwithstanding these limitations, our results are sufficient for
munities in our study. It was positively correlated with all an initial comparison with ground-level sites sampled in the
measures of the assemblage–total abundance, ordinal richness, same way. We found relatively few invertebrates utilising the
richness of all families, and richness and abundance of pollina- green roofs and the resulting small dataset has restricted the
tor species (bees and hoverflies). This finding is consistent with explanatory power of some of our analyses. For example, we
other studies which have found the abundance and/or richness have not been able to run multivariate mixed models that could
of bees (Tonietto et al. 2011; Braaker et al. 2014; MacIvor 2016; better elucidate the drivers of invertebrate responses. Similarly,
Ksiazek-Mikenas 2018), beetles (Braaker et al. 2014; Ksiazek- the level of taxonomic resolution we have been able to achieve
Mikenas 2018; Kyrö et al. 2018) and weevils (Braaker et al. 2014) limits interpretation of our results. Rather than focussing on a
increased with greater proportions of green space in the sur- specific taxa inhabiting green roofs and identifying samples to
rounding landscape. species level, because there have been no prior published stud-
The age and size of the green roofs did not consistently af- ies of invertebrates on Australian green roofs, we chose to ex-
fect the green roof invertebrate communities. Site age was amine the broad range of invertebrate orders occurring on
expected to increase habitat complexity leading to an enhance- them. The most abundant orders were further identified to fam-
ment of invertebrate taxa richness, similar to findings from ily level but resources did not allow species-level identification
ground-level urban green space (Smith et al. 2000; Sattler et al. which is problematic for many Australian insects due to the
2010), and a recent study in Melbourne that demonstrated that large number of undescribed species (Yeates, Harvey, and
older green spaces with more native vegetation cover provided Austin 2003). However, higher levels of taxonomic resolution
habitat for a range of native bee species (Threlfall et al. 2015). can reflect patterns at lower taxonomic resolution in terrestrial
However, green roofs are relatively new in Australian cities invertebrate communities (Timms et al. 2013).
Green roof and ground-level invertebrate communities | 7
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