Informal urban green space – A trilingual systematic review of its role for
biodiversity and trends in the literature
Authors’ manuscript, accepted at Urban Forestry & Urban Greening July 11, 2015.
Version of record: http://dx.doi.org/10.1016/j.landurbplan.2015.07.003
Citation: Rupprecht, C.D.D., Byrne, J.A., Garden, J.G., Hero, J.-M., (in press). Informal urban
green space: A trilingual systematic review of its role for biodiversity and trends in the literature.
Urban Forestry & Urban Greening. doi:10.1016/j.ufug.2015.08.009
Christoph D. D. RUPPRECHT1,2 (Corresponding author)
Address: Room 3.16, Building G31, Griffith University QLD 4222, Australia
Phone: +61 7 5552-9340
Jason A. BYRNE1,2 (firstname.lastname@example.org)
Jenni G. GARDEN1,3 (email@example.com)
Jean-Marc HERO1,2 (firstname.lastname@example.org)
1. Environmental Futures Research Institute, Griffith University, Nathan QLD 4111, Australia
2. Griffith School of Environment, Griffith University, Gold Coast QLD 4222, Australia
3. Seed Consulting Services, 106 Gilles Street, Adelaide 5000, South Australia
Urban greenspaces harbor considerable biodiversity. Such areas include spontaneously vegetated
spaces such as such as brownfields, street or railway verges and vacant lots. While these spaces
may contribute to urban conservation, their informal and liminal nature poses a challenge for
reviewing what we know about their value for biodiversity. The relevant literature lacks a common
terminology. This paper applied a formal definition and typology of informal urban greenspace
(IGS) to identify and systematically review a total of 174 peer-reviewed papers in English (152),
German (14) and Japanese (8). We identified three main topics: value for conservation (94 papers),
factors influencing diversity (80), and non-indigenous species (37). Additionally, we analyzed this
literature for temporal trends, spatial patterns, studied IGS types, taxa, climate zones, human impact
types, and key authors. Results show IGS plays an important role for biodiversity. Management
practices were identified as the most common and negative impact on diversity, while vegetation,
site age, distance to city center, and habitat diversity were positive-influence factors. The number
and impact of non-indigenous species varied widely. The analysis of literature patterns reveals: an
increase in publications over the last 15 years and a strong geographic bias in publications, as well
as towards temperate and humid climate zones. Studies of gap, powerline and microsite IGS were
scarce, as were studies of mammals and reptiles. Results suggest different maintenance regimes for
IGS may improve its contribution to urban conservation. We therefore propose adapting
management to the local context. (243/250 words)
Keywords: urban ecology; conservation; wasteland; spontaneous vegetation; cities, liminal
Some of the biggest conservation challenges, and most permanent ecological changes occur in cities
and towns (Goddard et al., 2010; Kowarik, 2011). Much of the research on urban forestry and urban
greening is dedicated to two types of spaces: (1) naturally vegetated spaces (e.g. remnants of the
pre-development vegetation), and (2) highly managed spaces with planted vegetation (e.g. formal
parks and gardens). Yet many scholars have emphasized the potential of spontaneously vegetated
spaces (e.g. brownfields, street or railway verges etc.) for urban conservation (Del Tredici, 2010a;
Kowarik, 2011; Kühn, 2006). For example, recent reviews concluded urban wasteland can
contribute to biodiversity conservation in urban regions (Bonthoux et al., 2014; Gardiner et al.,
2013), and quantitative research suggests such spaces cover around five percent of surveyed cities
(Rupprecht and Byrne, 2014a). However, knowledge of this topic is still quite limited. Most of what
we know is derived from English language literature. In contrast to research on parks and
conservation areas, research on informal green spaces also faces a conceptual challenge that
complicates identifying relevant papers – namely the lack of an agreed approach about how to
define these spaces.
In absence of a formal definition, researchers from urban geography and other fields have explored
the characteristics of informal green spaces. They argue such spaces are ‘liminal’ (Rupprecht and
Byrne, 2014b), and hard to identify and analyze because they form an ‘ambivalent landscape’
(Jorgensen and Tylecote, 2007) where land tenure, conservation, maintenance regimes, use,
regulation, and legitimacy are fraught with uncertainty (McLain et al., 2014). Liminality is a term
emerging from the social sciences (Rupprecht and Byrne, 2014a). It refers to a condition of
becoming, a transitional state of ‘in-between-ness’ or hybridity – distinguished by temporal and
spatial flux – and not easily categorized (Sweeney, 2009). As Pritchard and Morgan (2006, 764-65)
note, liminal spaces: ‘are borderlands between the mundane and the extraordinary…betwixt
places…[that are] mutable’. Head and Muir (2006, 506) assert that in liminal spaces can be found
‘complex entanglements of humans and nature…[where] …nature and culture are reinforced,
maintained or ruptured’ and ‘belonging is highly contingent’. Instone and Sweeney (2014) astutely
observe that for liminal ecologies, the culture/nature boundary is disrupted and divisions between
public/private and controlled/neglected are blurred. In sum, liminal spaces are ‘interfaces’ or
intersections of cooperation and competition, separation and reintegration, characterized by
informality and emergence (Imai, 2013).
The liminality of IGS may explain why researchers have referred to it using a variety of different
names, such as ‘urban wilderness’, ‘urban wildscapes’, ‘ambivalent landscapes’ or ‘urban
wasteland’ (Rupprecht and Byrne, 2014b). Without clearly specifying the object of study,
researchers risk overlooking important details about the attributes of these spaces and may remain
ignorant about a body of relevant and important previous research. Moreover, without definitional
certainty - that we are studying the same object, efforts to compare between different research
findings and to build knowledge are severely impeded. To address this issue, Rupprecht and Byrne
advanced a definition and typology of ‘informal urban green space’ (IGS) in a field survey of IGS
quantity (2014a) and provided a review of IGS’ role and value for urban residents (2014b). But
there is still a lack of knowledge about the biodiversity value of these spaces. This paper reviews
the scholarly literature on IGS and urban biodiversity, using the analytical framework provided by
Rupprecht and Byrne (2014a), offering researchers, planners, and stakeholders an integrated
understanding and synthesis of research findings.
Specifically, the review aims to address two sets of questions. The first set targets the role of IGS
for urban biodiversity: (1.a) how is IGS valuable to urban biodiversity conservation; (1.b) what
factors influence IGS biodiversity; and (1.c) how is IGS used by indigenous and non-indigenous
species? The second set of questions targets patterns and trends in the scholarly knowledge of IGS
biodiversity: (2.a) how has the number of relevant publications changed over time; (2.b) what is the
spatial and linguistic structure of the literature; (2.c) which IGS types have been studied most; (2.d)
which species groups have been studied most; (2.e) what forms of human impact are most common;
(2.f) what are the most studied climate zones; and (2.g) who are the key authors? These questions
assist in identifying knowledge gaps and identifying directions for future research. To answer these
questions, this paper provides a concise, tri-lingual review of 174 peer-reviewed research papers on
the biodiversity of IGS. Findings have important policy implications for biodiversity conservation
in urban areas.
We used a systematic review approach (Pickering and Byrne, 2013) that differs from a classic meta-
analysis. The systematic review has recently emerged as a useful tool for scholarly literature
analysis (Byrne and Portanger, 2014; Guitart et al., 2012; Roy et al., 2012). Such reviews do not
analyze published data; rather they identify geographic, theoretical and methodological gaps by
analyzing trends in the literature. Similar to a recent systematic review of the role of IGS for urban
residents (Rupprecht and Byrne, 2014b), this review included German, Japanese and English papers
to extend the scope of the review. These languages were chosen based on the multi-lingual
proficiency of the review’s first author. Preliminary searches revealed IGS-related research papers
published in other languages, such as Spanish (Lopez-Moreno et al., 2003) and Russian (Tikhonova
et al., 2002), and we recognize that we have not been able to address papers published in many
other languages (e.g. Mandarin, French, Portuguese etc.) – a point we return to in the discussion.
For this review, we systematically searched five major databases (Web of Knowledge, Scopus,
Google Scholar, CiNii and J-STAGE) using Boolean functions to combine search terms, for
example “urban AND species AND [all biodiversity terms with OR functions] AND [IGSvariable]”
(for full list of search terms in all three languages see Appendix A). Database searches were
performed in early 2011 for the full time frames available, and updated in early 2013 and late 2014
with a repeated search in Web of Knowledge, Scopus, Google Scholar, and J-STAGE for papers
published since the first search. We did not seek to impose a time limit on the search (e.g. 20 years)
but it should be noted that not all older papers may be full-text searchable, a limitation that may
cause them to be underrepresented. We selected a number of research papers specifically targeting
IGS to look in their reference sections for additional potentially relevant publications not returned
in the database searches.
To be included for analysis, publications had to meet three inclusion criteria: (1) the studied area
comprised or included at least one type of IGS following Rupprecht and Byrne’s typology (2014a,
2014b)(Table 1, Fig. 1); (2) the study reported sufficient details to identify a space as IGS (e.g. in
urban area, management arrangements, official park designation, site history); (3) the data reported
for an IGS was sufficient to include the study in the analysis of literature trends (e.g. target species
group). All feasible effort was made to clarify whether a study area fulfilled the requirements to be
included; aside from a close examination of all information provided in the publication, study areas
were (if possible) also located in Google Earth. Aerial photography and photographic material in
Google Earth was sighted to examine whether site conditions and site context in the urban matrix
complied with the three selection criteria above (a form of “ground-truthing”).
Figure 1 Photographs of informal greenspace types following the typology presented in Table 1. a)
Street verge, covered in spontaneous herbal vegetation (Brisbane, Australia); b) Lot, formerly
residential with perfunctory access restriction (Tōkyō, Japan), c) Gap, space between three
buildings with spont. herbal vegetation used by birds (Sapporo, Japan); d) Railway, annual grass
verge between rail track and street; e) Brownfield, spont. vegetated industrial space around
abandoned factory (Brisbane); f) Waterside, spont. vegetation on banks and deposits in highly
modified river (Nagoya, Japan); g) Structural, spont. vegetation growing out of vertical, porous
retaining wall (Tōkyō); h) Microsite, grass growing spont. growing out of crack in the pavement
(Nagoya); i) Powerline, vegetated right of way underneath high voltage powerline (Brisbane);
(Rupprecht & Byrne, 2014a).
Table 1 Informal urban greenspace typology (modified from Rupprecht & Byrne, 2014a)
trails and footpaths
Vegetated area within 5m from street not in another IGS category; mostly
maintained to prevent high and dense vegetation growth other than street
trees; public access unrestricted, use restricted.
Regular vegetation removal (>= once
per month); governmental and private
Soil, gravel, stone,
Vegetated lot presently not used for residential or commercial purposes; if
maintained, usually vegetation removed to ground cover; public access and
Irregular veg. removal, medium to
long removal intervals; private
Soil, gravel, bricks
Gap between walls
Vegetated area between two walls, fences or at their base; maintenance can
be absent or intense; public access and use often restricted.
Irregular veg. removal; variable
removal intervals; private
Rail tracks, verges,
Vegetated area within 10m adjacent to railway tracks not in another IGS
category; usually herbicide maintenance to prevent vegetation
encroachment on tracks; public access and use mostly restricted.
Regular veg. removal (monthly to
yearly); corporate or governmental
Soil, gravel, stone
Vegetated area presently not used for industrial or commercial purposes;
usually no or very infrequent vegetation removal and maintenance; public
access and use mostly restricted.
Irregular veg. removal, long removal
intervals; corporate and governmental
Vegetated area within 10m of water body not in another IGS category;
occasional removal of vegetation to maintain flood protection and structural
integrity; public access and use often possible with some restrictions.
Irregular veg. removal,
long removal intervals; governmental
Overgrown human artifacts; often vertical; occasional removal of vegetation
to maintain structural integrity; public access and use mostly restricted.
Irregular veg. removal,
medium to long removal intervals;
Soil, stone, gravel,
cracks or holes
Vegetation assemblages in cracks, may develop into structural IGS;
maintenance can be absent or intense
Irregular veg. removal, variable
removal intervals; variable
Powerline rights of
Vegetated corridor under and within 25m of powerlines not in another IGS
category; vegetation removed periodically to prevent high growth; public
access and use mostly unrestricted.
Regular veg. removal (less than
utility or governmental stewardship
Publications were systematically analyzed for findings on the role of IGS for urban biodiversity,
characteristics of each published study (year of publication, location, Köppen-Geiger climate type,
IGS description, target species group, species number or range found (where available) and human
impact). We also analyzed publication patterns across all research papers, such as temporal trends,
spatial patterns, studied IGS types, taxa, climate types, human impact types, and key authors.
Results are presented in tables and figures to efficiently present and synthesize findings from the
large number of articles, following similar presentation and analysis methods used in recent
literature reviews (e.g., Garden et al., 2006). Analysis of distribution among different climate zones
followed an updated version of the Köppen-Geiger system (Kottek et al., 2006) using a KMZ-file
(Wilkerson and Wilkerson, 2010). Principal and co-authorship was used to identify key authors who
contributed multiple articles.
We found a total of 174 papers, consisting of 172 original journal articles widely distributed across
90 journals, one book chapter and one Masters’ thesis. Journals publishing the most research papers
were Urban Ecosystems, followed by Landscape and Urban Planning, Diversity and Distributions,
Biological Conservation, then Journal of the Japanese Institute of Landscape Architecture (Table
2). This demonstrates that a variety of journals and scholars share an interest in this topic.
Table 2 Journals containing most papers on IGS biodiversity
Journals containing two or more papers
Number of papers
Percent of papers*
Landscape and Urban Planning
Diversity and Distributions
Journal of the Japanese Institute of Landscape
* Percentage does not add up to 100% as only journals with >3 papers are shown
3.1 Role of IGS for urban biodiversity
Research papers focused on three main topics: (a) value of IGS for conservation (94 papers), (b)
factors influencing IGS biodiversity (80), and (c) non-indigenous species found in IGS (37). A table
shows a summary of findings for the individual papers, including their publication year, location,
IGS type, climate zone, a detailed IGS description, details regarding human impact, the target
species group, number of species found (if available), and noteworthy comments about IGS and its
value (Appendix B). We discuss the main findings and their implications after summarizing the
results and examining trends in the literature.
3.1.(a) Value of IGS for conservation
The value of IGS for conservation was emphasized by just over half the papers (53%). Researchers
reported high species numbers across different IGS types and taxa (e.g., Brandes, 2001; Geibert,
1980; Muratet et al., 2007; Tan, 2010). Some IGS harbors rare species (Dana, 2002; Eyre et al.,
2003; Gilbert, 1990; Kadas, 2006) and was thus characterized as a wildlife refuge (Kantsa et al.,
2013). The contribution of IGS to biodiversity was often assessed in comparison to other areas and
habitats. Urban IGS can have higher species richness or diversity than rural areas (Mason et al.,
2006; Meek et al., 2010; Ray and George, 2009), lawns and forest (Robinson and Lundholm, 2012),
or ornamental plantings (Fründ et al., 1988; Vakhlamova et al., 2014), although non-indigenous
species may account for the difference (Ray and George, 2009). IGS can provide valuable habitat
(Brandes, 1992; Brown and Sawyer, 2012; Colla and Willis, 2009; Dallimer et al., 2012b; Rebele,
1988; Winter, 2013), and occasionally serve as a substitute for natural habitats (Joger, 1988; Kaupp
et al., 2004). It also represents an opportunity for urban residents to experience nature as a ‘natural-
cum-cultural’ heritage (Jim and Chen, 2011, 2010, 2008) or as a source of edible plants (e.g. in
urban foraging) (Diaz-Betancourt et al., 1999; Rapoport et al., 1995). While IGS can have
additional benefits for residents, this topic has been covered in our earlier review (Rupprecht and
Byrne, 2014b). We will return to how and why IGS can provide habitat and other benefits in the
3.1.(b) Factors influencing IGS biodiversity
A wide variety of factors influencing IGS biodiversity were identified in the research papers.
Scholars most commonly cited management practices and their negative impact on diversity (e.g.,
Helden and Leather, 2004; Jantunen et al., 2006; Jim and Chen, 2010; Vakhlamova et al., 2014),
even though habitat value for some indigenous species may depend on such management (Nemec et
al., 2011). Less direct disturbance may contribute to higher species numbers (Dana, 2002; Schadek
et al., 2008) by preserving vegetation communities valuable for conservation (Lenzin et al., 2007).
Different aspects of vegetation were regarded as important, especially vegetation structure
(Fernandez-Juricic, 2000; Florencia Carballido et al., 2011; Geibert, 1980; Strauss and Biedermann,
2006), vegetation as a food source (Eremeeva and Sushchev, 2005; Kazemi et al., 2011; Small et
al., 2006; Tommasi et al., 2004), and vegetation (including tree) cover (Ichinose, 2006; Itagawa et
al., 2010; Luther et al., 2008; Pennington et al., 2008). Biodiversity was found to increase with site
age (Crowe, 1979; Jantunen et al., 2006; Kim and Lee, 2005), distance from the city center
(Vakhlamova et al., 2014; Wahlbrink and Zucchi, 1994; Zorenko, 2003), and habitat diversity
(Dallimer et al., 2012b; Murgui, 2009), while it was negatively affected by sealed site surface (e.g.,
hard surfaces such as asphalt that can impede seedling growth) and substrate (Dallimer et al.,
2012b; Francis and Hoggart, 2008; Godefroid et al., 2007).
3.1.(c) Non-indigenous species found in IGS
Many researchers reported that they found high numbers of non-indigenous species across different
IGS types (Bigirimana et al., 2011; Garcillán et al., 2009; Kim et al., 2004; Ray and George, 2009),
particularly in New Zealand (Asmus and Rapson, 2014; de Neef et al., 2008), China (Gong et al.,
2013; Zhao et al., 2009), and the USA (Pennington et al., 2010; Stylinski and Allen, 1999). This
finding contrasts with papers reporting low numbers of such species (Catterall et al., 2010),
particularly in South-Africa (Cilliers and Bredenkamp, 2000, 1999) and Europe (Bornkamm, 2007;
Celesti-Grapow and Blasi, 1998). While some researchers reported that non-indigenous species
dominated (Asmus and Rapson, 2014; Crawford, 1979; Gantes et al., 2014; Stylinski and Allen,
1999), others found little evidence for competition (Celesti-Grapow et al., 2006). Some researchers
asserted that naturalized species may enhance urban biodiversity (Zerbe et al., 2004), provide
ecosystem services (Meek et al., 2010), and are of socio-cultural significance as they may possess
various desirable ecological and aesthetic qualities (Chmaitelly et al., 2009). Non-indigenous
species composition may also be used to trace historical patterns of introduction (Dehnen-Schmutz,
2004). While railway IGS was found to function as a corridor for grassland plants, it was not found
to provide any bonus to invasive species (Penone et al., 2012).
3.2. Trends and patterns in the literature
3.2.(a) Temporal trends
The earliest study included in our review was published in the 1960s (Bornkamm, 1961). Earlier
studies not appearing in our systematic search were reported in a post-war botanical study of
bombed cities (Lachmund, 2003). Over the last 15 years, the number of publications on IGS and
urban biodiversity has risen, with 70% of all research papers published since 2004 (Fig. 2). This
increasing interest could be related to ongoing global urbanization, the rise of urban ecology
(Douglas and Goode, 2011), as well as increasing recognition of the interconnections between
biodiversity and the well-being of urban residents (Dallimer et al., 2012a; Dearborn and Kark,
2010; Keniger et al., 2013).
Figure 2 Publication history of papers on IGS biodiversity
3.2.(b) Spatial and linguistic patterns
The geographic distribution of study locations in single-country papers shows a heavy bias towards
four countries: Germany (23 papers, 13%), the UK (22 papers, 13%), the US (18 papers, 10%), and
Japan (15 papers, 9%) (Fig. 3). Few research papers compared IGS in different geographical
contexts, causing a geographic concentration of knowledge about IGS especially in Europe (Fig. 4).
Papers from countries with increasing research output, such as China, are rare – a result possibly
caused by our limited capacity to search other languages, which we discuss in more detail later.
Research papers written in German (14 papers, 8%) and Japanese (eight papers, 5%) made up 13%
of all papers. Three German language papers studied IGS in Switzerland, while another one
compared IGS in multiple countries.
Figure 3 Geographic and linguistic distribution of papers on IGS biodiversity
Figure 4 Map of IGS biodiversity studies per country (including multi-national studies)
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3.2.(c) IGS types studied
Research papers that targeted at least two different types of IGS accounted for a third of all papers
(61 papers, 35%, Fig. 5). Brownfield and waterside were the most commonly studied IGS types in
single-type studies (27 papers or 16% each), followed by verges (22 papers, 13%) and structural
IGS (17 papers, 10%). Gap, powerline and microsite IGS were almost completely absent from the
literature. While some articles compared between types (Brandes, 2001), the number of IGS types
included in most multi-IGS-type papers was limited, which in turn limited potential comparisons.
As mentioned above, different authors may also refer to similar spaces by different names (e.g.
wasteland, derelict land, abandoned lot, vacant lot), which may complicate drawing upon their data
for potential future meta-analyses.
Figure 5 Distribution of papers on IGS biodiversity by studied IGS type
3.2.(d) Species groups studied
Vegetation dominated as the target of IGS biodiversity papers. Papers examining vegetation in
general were most common (79 papers, 45%, Fig. 6), but researchers also studied various subsets of
vegetation, such as vascular plants (5 papers, 3%), and groups of species not identical with a
specific taxon, such as spontaneous or non-native vegetation (4 papers or 2% each) or edible weeds
(2 papers, 1%). With regard to animals, birds (24 papers, 14%) and beetles (9 papers, 5%) were
most frequently studied.
Figure 6 Distribution of papers on IGS biodiversity by studied species group
3.2.(e) Human impact
Researchers have found a variety of anthropogenic influence types affect IGS. The most commonly
mentioned types were the design of the site and general maintenance/management (29 papers, 17%,
Fig 7.), followed by vegetation removal in the form of mowing, cutting or weeding (26 papers,
15%) and pollution of various kinds (24 papers, 14%). Aspects of site design such as substrate type
(e.g., bricks, gravel) were emphasized as particularly important for waterside (Francis and Hoggart,
2012) and structural IGS (Jim and Chen, 2011).
Figure 7 Most commonly mentioned types of human impact on IGS
3.2.(f) Climate zone distribution
Research papers showed a strong bias towards warm, temperate, and fully humid climate zones,
particularly Köppen-Geiger climate type Cfb (79 papers, 45%, Fig. 8), followed by Cfa (29 papers,
17%) and Dfb (18 papers, 10%). This bias likely results from the biased geographic distribution of
IGS biodiversity research sites and/or researchers (i.e. North America, Europe, and Japan).
Figure 8 Distribution of papers on IGS biodiversity by Köppen-Geiger climate zone
3.2.(g) Key authors
Five scholars contributed four or more of the research papers reviewed. Petr Pyšek analyzed trends
in urban vegetation diversity and composition over three decades (Pyšek et al., 2004) and co-
authored several papers on European IGS vegetation (Celesti-Grapow et al., 2006; Prach et al.,
2014; Prach and Pyšek, 2001; Pyšek et al., 2003). Cilliers and Bredenkamp studied the ruderal
vegetation of railway reserves, vacant lots, and road verges of South Africa (Cilliers and
Bredenkamp 1998, 1999a, 1999b, 2000). Brandes worked on ruderal vegetation of railway stations,
walls, and that of a small town (Brandes, 2001, 1992, 1983; Oppermann and Brandes, 1993).
Francis (with Hoggart) examined river walls and the influence of substrate on vegetation (Francis,
2011; Francis and Hoggart, 2009, 2008; Hoggart et al., 2012). Ten scholars contributed three
research papers as authors or co-authors, including Bornkamm (Abd El-Ghani et al., 2011;
Bornkamm, 2007, 1961), Jim and Chen (Jim and Chen, 2011, 2010, 2008), Kim (Kim, 2013; Kim
et al., 2004; Kim and Lee, 2005), Kowarik (Weber et al., 2014; Westermann et al., 2011; Zerbe et
al., 2004), Muratet (Maurel et al., 2010; Muratet et al., 2008, 2007), Pennington (Pennington et al.,
2010, 2008; Pennington and Blair, 2011), and Small (Angold et al., 2006; Small et al., 2006; Small
and Sadler, 2003). Twenty-six scholars contributed two research papers as authors or co-authors.
4.1 Role of IGS for biodiversity
Researchers have found that IGS plays an important role for urban biodiversity because it provides
a range of species with valuable habitat, as our systematic review of 174 research papers has shown.
This result is consistent with an earlier review by Bonthoux and colleagues (2014), who analyzed
37 papers and reported that the diverse local features of wasteland encourage diverse communities.
Our results further emphasize that the value of IGS depends on its local context.
IGS can provide habitat of a specific type otherwise scarce or absent in an urban area, for example
as structural IGS in the form of vegetated brick walls (Brandes, 1992). It may also resemble
ecosystems that were once dominant, but have declined as a result of landscape changes, such as
verges and brownfields with characteristics similar to sand plain grassland (Brown and Sawyer,
2012). By providing stepping-stones that support dispersal in urban areas, informal greenspaces
form part of a habitat network and enhance sustainability of metapopulations, as Kaupp and
colleagues (2004) reported for beetles nesting on spontaneously vegetated roofs. In addition to such
direct contributions to conservation, the localized socio-ecological aspects of IGS can produce
indirect benefits. In Hong Kong, spontaneous strangler figs may inspire awe in the viewer (Jim and
Chen, 2011), thus inspiring ecological awareness, and increasing the possibility of support for
nature conservation initiatives (Dunn et al, 2006). In Bariloche (Argentina), where malnutrition
poses a serious problem, 1.3 tons of edible weeds may be harvested per hectare of vacant urban and
suburban lots (Diaz-Betancourt et al., 1999), thus reducing the use of protected areas for
unsustainable livelihood practices. Such socio-ecological aspects can be important for biodiversity
because urban residents’ contact with nature likely influences conservation efforts beyond the local
urban area (Dunn et al., 2006; Millard, 2010; Miller, 2005). However, the main value of IGS for
conservation remains context-specific (e.g., which species benefit the most, and which likely do
not? Which type of IGS may provide which kind of threatened habitat type?). As Bonthoux and
colleagues (2014) argued, wastelands are not a uniform environment. The same is true for IGS,
which means planners and environmental managers must depend on localized knowledge to
effectively integrate IGS into urban conservation strategies – a point we return to shortly.
Factors influencing the biodiversity of IGS are characterized by two aspects, (i) the importance of
local features and (ii) the strong impact of management practices. Regarding the importance of local
features, our results are consistent with the findings of Bonthoux and colleagues (2014). Our results
further emphasize the importance of vegetation structure, vegetation as a food source, vegetation
cover, site age, and soil – in other words, characteristics that require planners to have a thorough
understanding of the local conditions in order to adopt appropriate conservation strategies (as
discussed above). The impact of management on IGS biodiversity was a widely reported issue in
the papers we reviewed, but was in contrast not reviewed by Bonthoux and colleagues (2014).
The expanded scope of our review casts a new light on the importance of maintenance practices and
their negative impact on diversity (Cilliers and Bredenkamp, 1998; Helden and Leather, 2004;
Jantunen et al., 2006; Jim and Chen, 2010; Namba et al., 2010; Vakhlamova et al., 2014; Yamato et
al., 2004). IGS, according to the definition used in this review (Rupprecht and Byrne, 2014a,
2014b), is neither formally recognized, nor its vegetation managed by its owner for agriculture,
forestry, gardening, or recreation. Yet various forms of maintenance (e.g., mowing, herbicide
spraying) are still regularly carried out (see above). Maintenance generally reduces vegetation
structure and complexity, in turn limiting the amount of food and shelter IGS can provide. This may
benefit pioneer and opportunistic species, but could make IGS less valuable for specialists. Some
maintenance may be necessary for utilizing the space (e.g., keeping verge vegetation from blocking
motorists’ line of sight (Brown and Sawyer, 2012)). However, as Hard (2001) pointed out, both
conservation-related and formal vegetation management in cities is ecologically and functionally
flawed: spontaneous vegetation is ‘managed’ using high levels of money, labor and herbicides to
protect abstract notions of aesthetics or risk minimization. Research by Nassauer has demonstrated
how aesthetics and social norms are important drivers for vegetation management (1988; 1992;
Nassauer et al., 2009), and as a result a perceived absence of management may signal a lack of care
(Nassauer, 1988), with flow-on impacts for biodiversity.
Such socially constructed ideals of greenspace (Lossau and Winter, 2011) and the notion that cities
are devoid of nature (long since dispelled by urban ecologists) may be reasons why IGS is often
viewed negatively and associated with decline (Corbin, 2003; Rall and Haase, 2011). To unlock the
potential of IGS to contribute to specific conservation goals, we may need to adapt management
practices accordingly. Brown and Sawyer (2012) provide examples for such adaptations in the
management of roadsides resembling sand plain grassland: changing mowing regimes to allow the
grasses to flower and mature seed could enhance the presence of rare species, while adjustments to
mowing height and width aid perennial species. This example demonstrates that management
adaptation is an intricate process. For such adaptions to succeed, we need to understand local IGS
conditions as well as the requirements of the species we aim to conserve.
Rare indigenous species have been found in IGS (Dana, 2002; Eyre et al., 2003), but so have non-
indigenous and invasive species (Asmus and Rapson, 2014) – an aspect that affects IGS
biodiversity management. Urban areas are characterized by challenging environmental conditions
that not all species are able to tolerate. While modified maintenance regimes may increase the
number of threatened species in IGS, even non-indigenous species that can adapt well to urban
environments may enhance biodiversity or provide ecosystem services. For example, Zerbe and
colleagues (2004) reported that non-indigenous vascular plants in industrial, road and railway sites
contribute close to a third of urban plant biodiversity in Chonju, South Korea. Moreover, Meek and
colleagues (2010) drew upon the concept of ‘novel ecosystems’ (Hobbs et al., 2006) to argue that
where restoration to historic conditions is not feasible, management should make use of non-
indigenous species to provide ecosystem functions. Importantly, IGS does not replace formal green
space such as parks, gardens and conservation areas. Rather, IGS is a liminal, hybrid, socio-
ecological entity that provides habitat for plants and animals as well as opportunities for urban
residents to interact with and experience nature (Rupprecht et al., in press; Rupprecht et al., 2015;
Rupprecht and Byrne, 2014b). Therefore, researchers have suggested spontaneous vegetation could
be understood as the “de facto native vegetation of the city” (Del Tredici, 2010b) because it is
always appropriate to site conditions (Kühn, 2006). This affects policy recommendations, discussed
in more detail later.
4.2 Trends and patterns in the literature
Our results have revealed a strong bias in the reviewed IGS literature towards specific regions
(Europe, the USA, and Japan) and climate zones (temperate and humid such as Cfb, Cfa, and Dfb).
One limitation of our review was our capacity to search other languages besides English, German,
and Japanese. This limitation likely contributed to the spatial bias we found in the literature.
However, papers published in both German and Japanese only accounted for about half of the
studies conducted in Germany and Japan, even though the different linguistic distance between
English and the two languages (Chiswick and Miller, 2005) makes learning English easier for
German researchers than for Japanese researchers. This could suggest that the comparatively low
number of English publications on IGS biodiversity may not solely result from missing non-English
publications, but could instead indicate an actual gap in our knowledge about IGS biodiversity in
these countries. Future reviews should therefore target additional languages to clarify this issue.
If we lack local IGS knowledge, the spatial and climate zone bias is a major concern, because it
would impede our ability to devise context-specific conservation measures in regions that are home
to large urban populations, such as China, India, South-East Asia, Africa, and South-America. In
particular, climate zones A (four studies) and B (11 studies) are severely understudied, but account
for 88% of Africa, 75% of South America, and almost all of South-East Asia (Peel et al., 2007).
Countries in these regions are experiencing both rapid urbanization (UN-HABITAT, 2012) and
threatened biodiversity (Zhao et al., 2006). But it is possible that there is a literature on IGS in these
climatic zones that has not been explicitly framed around biodiversity conservation. For example, in
the megacities of Africa and Asia, there may be an emphasis on food security rather than
biodiversity. Urban interstices offer the potential for growing food, especially for socio-
economically marginalized and vulnerable populations, and for growing medicinal herbs. Growing
plants valued for their medicinal properties or nutritional benefits does not necessarily diminish
biodiversity, and recent studies of urban food gardens have shown that they can be highly
biodiverse (Galuzzi et al., 2010; Weinberger, 2013). Therefore, a better knowledge of local IGS
could help to devise strategies for preserving urban biodiversity in these areas, which depend on
local knowledge to be effective (see above).
Studies on brownfield, waterside, verges, and structural IGS types were the most common, while
gap, powerline and microsite IGS are still comparatively understudied. The area of these
understudied sites is usually much smaller than that of a vacant lot or brownfield IGS, which may
make such sites seem like a less rewarding object of study, and/or present significant
methodological challenges. However, the fragmented nature of urban landscapes makes it likely
that a high number of such spaces exist within cities. For example, a recent case study suggested
that almost 20% of IGS, or one percent of the surveyed area in Sapporo (Japan) consisted of gap
IGS (Rupprecht and Byrne, 2014a) – an amount particularly valuable for conservation in dense
urban areas where other greenspace is scarce. These hitherto little-examined IGS types also warrant
closer attention because different IGS types differ in their characteristics (Table 1), and may
consequently contribute to urban conservation in different ways. A better understanding of gap and
microsite IGS may also help planners to create synergies between conservation and greenspace
strategies. Specifically, they may be able to act as additional stepping-stones, similar to vegetated
roofs (Kaupp et al., 2004), while contributing to the prevention of urban heat-island effects.
Studies on the vegetation of IGS and its role for birds and beetles were comparatively common, but
we presently know little about if and how IGS can be valuable for mammals and reptiles. Studies on
ants were also scarce, despite research suggesting vacant lots can feature a distinct species
composition and can be richer in species than gardens (Uno et al., 2010). While the limited size of
some IGS sites suggest their value could be limited, large or linear sites such as powerline and
railway verge IGS could potentially function as movement corridors for large urban wildlife (e.g.,
coyotes, foxes, deer, kangaroos) connecting urban and peri-urban areas (Rudd et al., 2002).
A number of authors (e.g., Cilliers and Bredenkamp in South Africa, Jim and Chen in Hong Kong,
Kim in South Korea) that contributed three of more studies were based outside of Europe, the USA,
and Japan. This stands in contrast with the regional bias of the literature. Knowing authors central
to the field is important, because it allows us to understand how the current body of IGS literature
developed. Additionally, it provides a starting point for studies on the history of IGS biodiversity
science. Such authors possess valuable expertise that may help in devising locally adapted
conservation strategies. They could also play a role in coordinating future research efforts in their
regions, or collaborate for cross-regional and cross-cultural studies as follow-ups to emerging cross-
national studies (e.g., Lososová et al., 2011).
5.1 Policy recommendations
Our review of 174 research papers on the role of IGS for biodiversity found that IGS is valuable for
conservation, but appropriate management is important for maintaining IGS biodiversity (though
this must be inferred because few, if any, studies have demonstrated a statistically significant
correlation). We therefore propose to complement the suggestions for conservation and planning of
urban wastelands by Bonthoux and colleagues (2014) with a review of maintenance practices. For
example, reducing or changing mowing intervals may not only benefit site diversity (Brown and
Sawyer, 2012) and save resources, but may also preserve the natural site character that residents
cherish (Rupprecht and Byrne, 2014b). However, planners should avoid treating IGS like
conservation areas by restricting residents’ access, as the diversity of formal and informal uses
produces the habitat diversity and local features that make IGS valuable for biodiversity (Bonthoux
et al., 2014; Hard, 2001). A thorough understanding of these local features and the local context
should inform IGS management, and facilitate integration into urban conservation strategies.
Planners and government agencies need to work with owners of IGS, such as utilities and railway
operators, to phase out harmful maintenance practices (e.g., herbicide spraying). Where frequent
vegetation maintenance is essential or strongly preferred as a result of residents’ preferences
(Nassauer et al., 2009), encouraging a conversion of IGS toward recreational green space types such
as community gardens may be an option. For example, the power utility Chubu Electric Power
invites local residents in Nagoya (Japan) to use land under urban power transmission lines for
gardening free of charge, if they in return keep vegetation under a specified height (Rupprecht and
Byrne, 2015). The utility profits financially from reduced maintenance expenses, the community
enjoys additional recreational opportunities, and birds as well as insects gain a source of food. As
such arrangements in particular and the conservation value of IGS in general are determined by its
local context, we propose directions for future research to fill the gaps in our local knowledge of
5.2 Directions for future research
This review has identified three major gaps in our knowledge of IGS, our localized knowledge of
IGS around the world, our knowledge of understudied IGS types, and our knowledge of
understudied species groups. First, we know little about IGS biodiversity outside of the temperate
and humid Cfb, Cfa and Dfb climate zones of Germany, the UK, the USA, and Japan. Future
research should target IGS biodiversity in South-East Asia, Africa, South America, the Middle East,
India, China, and Australia, as well as IGS in the climate zones A and B. Moreover, international
comparisons of IGS are rare, and the lack of studies in many regions limits potential meta-analyses
and cross-cultural studies. How do different cultural contexts influence the value of IGS for
biodiversity, the possibilities for management adaptions, or the potential for hybrid conservation-
recreational use? However, it is important to note that this review only examined the available
literature in English, German and Japanese. As discussed above, our search also found Spanish and
Russian research papers on IGS. A review of literature on IGS in these languages, Chinese, French,
Indonesian, Polish and other languages would likely advance our understanding of IGS and help
local planners and IGS owners to adapt policies and management.
Second, we lack studies on gap, powerline and microsite IGS as well as comprehensive
comparative studies. Future research should address this lack of knowledge by examining some of
the following questions. How do gap, powerline or microsite IGS contribute to urban biodiversity?
How does their potential contribution compare to other IGS types? How can management practices
for these sites be adapted to benefit conservation? Moving to study designs based on a common IGS
typology may help us to identify urban habitats important to biodiversity that researchers might
have previously overlooked, and could facilitate studies comparing between different IGS types.
Research on smaller sites could also redress the paucity of knowledge about IGS in the megacities
of Africa and Asia. For instance, it might help to answer questions about whether IGS is meeting
food-security needs, such as the harvesting of spontaneous vegetation or the growing of ‘bush
foods’ in the urban interstices, and how in turn this might impact biodiversity.
Third, future studies should investigate the role IGS may play for hitherto scarcely studied species
groups. Can IGS benefit mammals, reptiles, or marsupials? Do limited size and human disturbance
prevent large animals from using IGS? How does the presence of animals in IGS affect resident
perception (e.g., opportunities for nature contact, potential for wildlife conflict)? We need to
address these three main gaps in our knowledge. Closing these gaps would be a first step to better
understanding the local features of IGS – local features that are key to how IGS contributes to
biodiversity, how we should adapt our management of IGS, and how we can integrate IGS into
urban conservation strategies. Better knowledge of IGS is crucial for future conservation efforts in
Finally, the increasing number of studies on IGS biodiversity provides a growing source of data that
future studies could draw upon for meta-analyses. For example, IGS size did not feature
prominently as a driving factor for species diversity in the papers we examined in our study –
despite the important role of this factor in ecological theory (e.g., island biogeography). Future
research could analyze a set of IGS studies to explore what role IGS size and related factors such as
fragmentation play for the biodiversity of these urban spaces. Another potential target for a meta-
analysis would be to quantify (using statistical analysis) the apparent negative relationship between
the degree of IGS management and IGS biodiversity – as suggested in some of the literature
addressed by this paper. We recognize that this is just one step in a much larger research agenda on
IGS. Future studies could address diverse aspects of this understudied component of urban forestry
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Appendix A – Search terms used in English, Japanese, and German
(michi no hashi)
(hodō no enseki)
trails, foot paths
Weg, Pfad, Fusspfad,
Appendix B (online suppl. info?) – Author, year, location, IGS type, species group, study area, climate zone, IGS description, species number,
human impact, and comments on IGS of all 174 individual research papers
Human impact on
Value and comments regarding IGS
Flora distinct from other urban habitats
Species diversity increases with aridity,
soil character changed by anthropogenic
Dispersal between sites important for
flora, chain of habitats, recommend
Separate seed pool from urban ecosystem,
Ruderal & waste areas,
walkways, walls, lawns
89% exotic species, town flora very
homogenous, environmental influence
factors include distance from coast & size
of central business district
No decay of compositional similarity with
increasing spatial or environmental
distance was found
Disturbed reach had lowest herpetofauna
abundance and species richness, increased
vegetation structural complexity
trampling, grazing, fire
High abundance of introduced species
Variety of plant communities, extreme
wet and dry conditions
Bare experimental plots
over 38 years
Alien species rare, results support
spontaneous succession as cheap way to
develop near-natural plant communities
rich in species
Human impact on
Value and comments regarding IGS
Active and abandoned
core rail yard areas
Abandoned rail yards of special
importance, valuable for biodiversity
Important as habitat and for biodiversity,
recommendations for plant-friendly
Stone and walls, verges,
riverbanks, rail tracks,
rail yard, wasteland
Highest diversity in wasteland and rail
Complex upland grassland habitat
reminiscent of agricultural grasslands in
19th century; not ecological wasteland
Riverbank (0-20m from
Presence, food waste,
Vegetation and veg. Diversity important
Road strip corridors
Can function as corridors, can contribute
to gamma diversity, potential not
recognized by authorities
Vacant lots, rubbish
dumps, stream banks,
railway banks, vacant
Food waste, shelter,
Health risk, examined spaces provide
Suburban road verges
Relatively high diversity and thus
valuable, may increase diversity if
replacing agriculture, homogenization not
supported, low replacement of natives by
Ruins, dumping sites,
industrial sites, road ides
Intense human use
Flora not uniform between cities, high
diversity, low alien diversity and
Archeological sites, new
wasteland and vacant
lots, historical center
with spon veg,
Intense human use
No competition between natives and
aliens, high diversity, diversity dependent
on habitat and disturbance
Human impact on
Value and comments regarding IGS
unmanaged soil, walls,
sidewalk, planted beds
Trampling, distance to
Microhabitats similar in species
composition, could be used to enhance
species diversity in city center
Diversity may have been decreased by
more frequent maintenance, increase of
ruderals and aliens due to increased
Road gap, abandoned
land (soil or gravel)
land use change,
Species diversity much lower than in
former non-IGS land use, increase in
xeric and mesic species
Birds exploited green walls but were
never found on bare walls, veg. walls can
provide resources for birds without
Vacant lots, coastal cliffs
High floral diversity, recognize that
naturalized flora have various ecological
as well as aesthetic qualities and socio-
Roadside green, bridge,
ruderal sites, waste
Anthropogenic habitats bear a poor and
apparently random proturan fauna - yet
contribute one sixth to the overall species
Low species number per sample plot in
comparison with natural areas,
management should encourage
Previously undescribed communities,
conservation not necessarily means
changes in maintenance practices
Disturbed soil (post-
Relatively low percentage of introduced
species (35%), no similarities with ruderal
communities in other continents
Well-established vegetation, low
percentage of introduced species (26%),
higher than similar ruderal sites in the city
(see Cilliers 1998, 1999a, 1999b)
Human impact on
Value and comments regarding IGS
Derelict demolition sites
Disturbed soil, brick
Cheap landscaping could increase
potential use, diversity and attractiveness
could be increased by sowing seed
collections from other wasteland sites
None after construction
Green roofs can offer habitat for a variety
of bee species
Former dumping site
with surface earth fill
Low arthropod diversity, absence of low
dispersal ability taxa, spider fauna
dominated by an introduced species
Diversity increases with age and lot size,
decrease with isolation
Important part of urban habitat mosaic,
influence of habitat diversity (positive)
and sealed surface (negative) on species
Vacant lots, walls,
of vegetation possible
several times a year
Should be considered for conservation,
contain rare species, balance between
protection and needed disturbance
Frequent spraying and
High number of exotic species, numerous
benefits of wall vegetation, great potential
(large area, additional vertical space for
densely developed districts)
Castle rocks and walls
Number of usable exotic plants show
historical reasons for introduction
Rare species excluded, up to 60% of
variance in spont. Plant distribution was
explained by pollutant dispersion pattern
43 (Coatepec), 32 (Bariloche)
Significant potential as food source
providing more than 1 ton per ha of edible
Minimally managed long
Vegetation more important for small
mammals than urban environment factors
Human impact on
Value and comments regarding IGS
vacant lots, railway and
Preservation important because vegetation
could form dispersal corridors
Landfill provides stable habitat, but
drainage facilities critically affect beetles
(fall into drainage)
Large areas of urban plots with partly
restored vegetation provide sufficient
food supply for butterflied and
bumblebees, pollution important for
Various brownfield sites
(railway, factory, canals)
Large number of rare species, high
Vegetation structure and park connection
have positive influence
Reduced vegetation due
to landfill legacy
Mostly indigenous species, can play role
in conservation, vegetation structure
factors explain most abundance data
Vegetated vacant lots
No similarity to other vacant lot studies,
many therophytes, usually one
dominating species per community
Maintenance, choice of
Strong influence of substrate material on
habitat potential, brick and boulders
preferred to concrete, conservation
choice of substrate
Mix of terrestrial and riparian species,
surface fractures increase plant diversity,
habitat improvement potential
"Mass effect" - flora maintained by
propagule pressure, significantly more
diversity on bricks than sheet metal,
potential for habitat improvement
Soil biota and
space, verges, street tree
High diversity, wasteland and verges
more diverse than flower plantings
Human impact on
Value and comments regarding IGS
Partly active landfills
Exotic species are dominant, natives gain
with age of cells, in oldest cells some
species belong to local climax community
High percentage (61%) of non-natives in
comparison to other vacant lot studies
Lower diversity than other urban
landscapes, but different microlandscape
types harbor different species
High diversity, higher than in neighboring
residential area, vegetation structure
complexity and cover over 60cm
correlated with bird diversity
Highly urban, recently
bonfires and children's
Higher than expected diversity, rare and
newly discovered species, threatened by
development and economic growth
Derelict and despoiled
Former land use,
Probability of species occurrence related
to land use
Former industrial area,
demolished house lots
Concrete substrate and walls around a site
lowered diversity, different anthropogenic
substrates have different flora
Linear corridors along
roads and sidewalks or
Verges similar to residential and
industrial vegetation in native-alien ratio,
alien species widespread
Abandoned ruderal area
Building density and use intensity
influence carabid distribution
Wall vegetation contributes to urban
diversity and to the visual character of the
city center and thus deserves protection,
human beauty perception plays a role in
Brick kiln brownfield
Brick and ash rubble
Varying diversity in different seasons,
less diversity due to brick dust stress
Human impact on
Value and comments regarding IGS
Urban ruderal communities may comprise
consistent and separate plant associations
roadside, empty lots
C3 and C4 alien species prefer ruderal
habitat compared to the native species
Riverbanks and islands
Elimination of dominant alien plant has
temporary positive effect on native plant
richness but causes other alien plant to
Road management practices favor
ephemeral annuals and short-lived taxa,
arable land weeds dominant
Roundabouts and other
Grassland Hemiptera diversity would be
increased with a reduction in the intensity
of management, such a reduction in the
frequency of mowing
Flood defense walls
Wall design choice
Highest richness on brick walls, lowest
richness on concrete walls, influence of
algal cover and river flows
Strong human influence
Different levels of anthropogenic
disturbance are reflected in the two rivers'
Urban matrix (perching
Strong relationship with vegetation cover
and >2ha woodlot vicinity
High diversity compared to other urban
Wooded streets on
Vegetation height, tree cover and distance
to original land are related to inhabitation
(drastic chemical and
Verges are distinct from semi-natural
grasslands, are species-poor due to young
age, over-management and disturbance
but show potential if these conditions
change (old, unmanaged verges)
Stone retaining walls
Precious ecological asset, natural-cum-
cultural heritage, threatened by misguided
Human impact on
Value and comments regarding IGS
Land use, wall
Ecological heritage, environmental and
visual amenities, need to be protected
Conservation and biodiversity value,
places of nature-in-city, beneficial win-
win situations possible
High diversity, may act as substitute for
disappearing natural habitats (cliffs)
High diversity of species, substrate,
structure and processes
High diversity and large future potential,
Old stonewall, rubble,
vacant lots, building
walls, fortress wall.
Plants of conservation interest present,
wildlife refuge character
High diversity, function as stepping stone
and natural habitat substitute
Lawn-type street verges
Monoculture lawn with intense
management and low biodiversity
Lawn-type street verges
Comparatively low diversity, negative
impact of missing flowering plants
Possible to support succession to typical
forests, comparatively high number of
Soil seed bank important, age related to
Waste landfill with
Fill materials, soil
Succession is a viable option for
restoration unless no nearby propagule
source is present
Areas serve variety of bird species
groups; influence of slope, artificial
structures and vegetation
Difference in urban and suburban sites,
influence of water quality, vegetation,
Human impact on
Value and comments regarding IGS
May have functioned as habitats under
regular mowing, can serve as key
reservoirs for recovery
Non-native species not discovered before
Wall tops, verticals
Substrate choice, air
High diversity, nutrient and moisture-rich,
mostly common species
Verges, roofs, cracks
Some neophytes resistant to urban
disturbance, but outcompeted by natives
in other places
Industrial area, harbor
High conservation value, absence of
anthropogenic disturbance causes
632, 675 (plants), 40, 73 (snails)
High diversity esp. in mid-successional
sites, high conservation value, endangered
Riparian surrounded by
Infrastructure, land use
Infrastructure and residential areas have
most influence, benefit tolerant species
Urban riparian areas
Main factors influencing diversity are tree
cover percent and shrub species richness
Abandoned lots, small
Abandoned lots have highest abundance,
offer better breeding conditions than
Provide habitat for high number of native
plants, "wild roof" as potential rooftop
Urban riparian areas
Diversity key influence is dominance by
invasive species (regardless of nativeness)
Urban riparian areas
Urban areas have higher species richness
than rural areas
Vacant urban land,
R. japonica negatively influences other
species, but covers not more than 4% per
Area provides rare open space habitat for
wild plants within Berlin
Human impact on
Value and comments regarding IGS
Urban riparian areas
Land use regime
Urban areas have higher species richness,
alien species can provide ecosystem
Value for endangered species, no impact
of human and dogs, greenspace design
Edges and center of
Increase of weeds without herbicide, but
not very pronounced
Lower species richness than any other
land use type
Disturbed river banks
Large number of ruderal species, soil
texture and topography strongest
Relatively high diversity, tall trees
recommended to attract tree-reliant
Areas with abandoned
Wasteland has highest species richness of
all habitat types, 20% naturalized species
Highest floristic interest index habitats
semi-natural, dwellings exhibits neg.
Built-up land cover
Positive influence of habitat diversity,
negative influence of built-up habitat
Verges, vacant areas
Population decline due to intensified
Urban trails along (e.g.)
Habitat value for native species may
depend on intensive management
High number of indigenous species, high
overall species number, important for
Brownfields, rail and
road verges, walls,
Conservation value of strongly
transformed habitats pose conservation
Ruderal, industrial or
Ruderal area has highest species richness
and density, high conservation value
Human impact on
Value and comments regarding IGS
Urban riparian areas
Canalized areas less diverse than un-built
ones, many neophytes but little use of
river as vector
Spontaneous woody vegetation plots had
higher bird diversity, plot size important
for plants and birds
29% of probable horticultural origin,
derelict railway walls have higher variety
Riparian edges in
Tree cover, native vegetation and building
area influence opposite for native and
Riparian edges in
Native species decrease, non-native
increase with urbanization, some natives
Riparian edges in
Habitat selection factors operate on both
proximate and broader spatial scales
Railway edges function as corridors for
common grassland plants but provide no
bonus to invasive species
Seasonal fluctuations of species richness
between urban/rural areas
Ruderal urban sites
Spontaneous succession can be relied
upon for restoration projects, cheap
Road verges, ruderal
urban sites, abandoned
Sere identify was not sign., sere
vegetation formed continuum along
moisture gradient and by successional
age, spontaneous succession mostly
results in woodland and is ecologically
suitable restoration option
Dump area, human density in region and
altitude positively influence species
Ruderal urban habitats
Change in construction
practice, winter salt use
Decrease in archaeophyte species richness
and diversity from 1960s to 1990s,
Human impact on
Value and comments regarding IGS
Road and railway
Corridors cover only 2.7% of city but
hold 76.3% of flora, CR-strategists
prevail, corridors resilient to disturbance
Disturbed suburban lots
Cultural preferences for
Edible weeds can provide considerable
food source, should be used to
Urban areas have higher species richness
than rural areas, more exotics
Use (industrial, kids),
Decrease of derelict areas leads to
dwindling wild flora habitats
Most species grow better on base of wall,
less diversity than in Europe
Higher plant species diversity,
invertebrate abundance and taxonomic
diversity than lawns and forest
Don river banks
River banks provide habitat to bird, plant,
butterfly and macroinvertebrate species,
benefit from river connectivity
Road kill, pollution,
Important reserve for some species,
diversity similar in different road verge
Ruderal areas, verges,
Relict areas form basis of rich species
composition, but threatened by
Rare plant species important in
determining rare Hemiptera species
Artificial coasts are colonized by plants
with floating seeds but not by those
abandoned railroad, new
Soil alteration (rubble,
High plant species richness possibly
achieved by strong disturbances every 5
Port, industrial sites,
railway system, traffic
Diversity similar between urbanization
zones, high number of species
Human impact on
Value and comments regarding IGS
Past herbicide use
Influence of surrounding gardens, areas
contribute to urban biodiversity
abandoned fields, refuse
areas, railway, roads
Fire, cutting, digging,
trampling, waste dump,
Urban vegetation favored disturbance,
nutrient and water resources are abundant
Disturbance by motor
Enrichment from adventitious species, but
species composition loss in technogenic
Former factory, housing
an railway ground
Most species rich assemblages found on
early successional sites
Habitat quality (early successional sites
with diversity of seed producing plants)
Channel discharge changes and riparian
vegetation changes controlled channel
146/130 (LH), 11/15 (GH)
Vegetation structure most important,
species prefer certain succession stages
disturbed sites (e.g.
military training ground)
Exotic species dominate, native species
cover low even after 70 years
Tramlines and building
Higher number of therophytes, many
(light) tree seedlings on building surface
Highly modified river
bed and banks
land use, water quality
Complexity of land use and
environmental quality affects birds,
ground beetles and plants
Vacant lot vegetated
High diversity despite small area and high
Bloom and habitat heterogeneity are key
to urban area potential for bees
Urban structure (both land use and
vegetation) best described potential
Former residential use
Exotic species abundance correlates with
ant species richness
Human impact on
Value and comments regarding IGS
land, landfills, eroded
deposit, fire, industrial
Species diversity increased with distance
to city center, species richness at
unmanaged sites higher than at
ornamental sites, alien species lowest
Matrix grassland on
Urban dry meadows important habitats,
but matrix grassland least diverse,
important to avoid replacement with
Low diversity per site but high
discrimination among lots
Towards city center shannon diversity,
evenness and carabid body size decrease
Not dedicated diversity survey, roadside
spont. Vegetation immobilizes significant
amount of air pollutants, increasing
biodiversity supports air filtration
Environmental and landscape predictors