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Urban Ecology and Special Features of Urban Ecosystems

Author(s): Franz Rebele


Source: Global Ecology and Biogeography Letters, Vol. 4, No. 6 (Nov., 1994), pp. 173-187
Published by: Wiley
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Global Ecology and Biogeography Letters (1994) 4, 173-187
A
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Urban ecology and special features of urban
ecosystems
F R A N Z R E B E L E Institute of Ecology, Technical University of Berlin, Schmidt-Ott-Str. 1,
D-12165 Berlin, Germany
Abstract. The paper deals with urban ecology as a
biological science and applies some of the topics of
general importance in ecology to the special condi-
tions found in towns and cities. I consider whether
cities should be treated as one integrated ecosystem,
or as an assemblage of various ecosystems. In contrast
to the holistic, organismic concept of the ecosystem as
a new hierarchical level of organization and as an
evolving whole which guides the development of the
species, I follow the methodological definition of
Tansley (1935), who defined ecosystems as 'mental
isolates' for 'the purpose of study'. According to
Evans (1956) ecosystems can be defined at every level
of the biological organization, at the level of the
organisms, populations or communities.
The introduction of species from other biogeo-
graphical regions is a worldwide phenomenon, but the
proportion of successfully established introduced spe-
cies is higher in cities than in rural or forest areas.
This is due to numerous colonizing species which fit
the anthropogenous habitats. Due to unequal rates of
immigration and extinction of species, urban habitats
show an imbalanced turnover of species.
Another special feature of urban ecology is man-in-
duced disturbance, which initiates the colonization of
disturbed or newly created habitats. According to the
type of substrate and the availability of diaspores
there may be both primary, secondary or intermediate
types of succession. Besides disturbance, the main
component for structuring communities is biological
interactions. In this paper I discuss some aspects of
competition, predation and mutualism.
The special feature of higher species' richness of
cities compared with ecosystems in the countryside
can be explained by the high habitat diversity of urban
and industrial areas. Although some components
which contribute to the complexity of communities,
such as competition, are of minor importance in vari-
ous urban habitats, there may be communities of high
complexity.
I also consider community characteristics such as
stability and productivity. Since most urban communi-
ties are in a state of inequilibrium, theories of stability
based on equilibrium are inadequate for urban ecosys-
tems. The productivity of the 'ecosystem city' mainly
depends on the area of unsealed open space and the
successional stage of the plant communities of the
various habitats.
Key words. Ecosystem city, biological invasions,
colonization, diversity, stability.
INTRODUCTION
Urban ecology has become an expanding field of
research during the last two decades. Various studies
were carried out on urban climate, soils, flora, fauna,
urban habitats and green space of cities (Berry &
Kasarda, 1977; Bomkamm, Lee & Seaward, 1982;
Duvigneaud & Denayer-de Smet, 1977; Exline, Peters
& Larkin, 1982; Gill & Bonnett, 1973; Gilbert, 1989;
Hollis, 1991; Kieran, 1982; Klausnitzer, 1993; Laurie,
1979; Sukopp, 1990; Sukopp & Hejny, 1990; Sukopp
& Wittig, 1993; Wittig, 1991; for reference see also
Sukopp & Werner, 1982; Sukopp et al., 1986; Dawe,
1990). Most of this work has been done to solve
practical problems of urban planning, environmental
protection and urban wildlife conservation. On the
other hand, theoretical and conceptual work in urban
ecology was hardly stressed (Trepl, 1994).
Urban ecology is both a practical science dealing
with the environment of people living in towns and
cities, and the associated 'environmental problems'
such as water, air and soil pollution, extrac-
173
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174 Franz Rebele
tion of drinking water, transport planning, noise, etc.,
and a biological science as well. As a sub-discipline
of ecology, urban ecology is concerned with the distri-
bution and abundance of plants and animals in towns
and cities. As with other ecological disciplines it is
possible to distinguish between organizational levels
of individual organisms, populations and communi-
ties.
The practical aspect of urban ecology is not dealt
with here. In the following, I discuss some ecological
concepts and principles as they apply to urban ecosys-
tems.
ECOSYSTEM CITY AND URBAN ECOSYSTEMS
A question often asked in urban ecology is whether
towns and cities can be viewed as 'ecosystems'. In
'The use and abuse of vegetational concepts and
terms', Tansley (1935) defines an ecosystem as a
system which takes account both of the organisms and
the entire complex of physical environmental factors.
Ecosystems are 'mental isolates', set up for 'the pur-
pose of study' (Tansley, 1935, p. 300). This is primar-
ily a methodological definition, and the limits of the
ecosystem are drawn by the scientists involved; it can
form both a part of larger systems, can overlap with
other ecosystems or can interact with these.
According to Evans (1956), the definition of
ecosystems as the basic unit in ecology does not mean
its definition as a new hierarchical level in the organi-
zation of communities. Ecosystems can be defined at
every level of biological organization, at the level of
the organisms, populations or communities, and he
recommends that the level be indicated by an appro-
priate combination (community ecosystem, population
ecosystem).
Begon, Harper & Townsend (1990) also argue that
no ecological system, whether individual, population
or community, can be studied in isolation from the
environment in which it exists. Therefore they do not
distinguish any separate ecosystem level of organiza-
tion. Domains of ecosystem researchers such as nutri-
ent dynamics and energetics are dealt with in
connection with the structure and function of com-
munities. This approach takes due account of the fact
that ecology as a science investigates the interactions
between organisms and their environment, and also of
the fact that an environment considered in isolation
from the organisms whose environment it fornms is an
empty concept.
In contrast to the view that communities or (com-
munity) ecosystems can be described by the sum of
their parts (i.e. the population of species) plus their
interactions, the ecologists of the holistic school see
ecosystems as an important hierarchical level of orga-
nization of life on earth (c.f. Odum, 1971; Pomeroy &
Alberts, 1988.)
This view of ecosystems is in the tradition of
Clements' organismic concept of the community
against which Tansley (1935) originally wrote.
Ecosystems are seen as evolving entities, which guide
the development of the species and which in principle
are independent of the species combination. These
ecosystems are attributed properties similar to those of
organisms, such as self-regulation. At the core of this
direction of ecosystem research is the energetics of
ecosystems. (On the discussion of this, see e.g. McIn-
tosh, 1980; O'Neill et al., 1986, Trepl, 1988).
Looking at the parameters discussed in ecosystem
research, there are none which cannot be explained at
the organizational level of the communities and popu-
lations forming them, e.g. species richness, diversity,
productivity, energetics of the system, stability, re-
silience). The level of 'self-regulation' (if any) can be
explained by the activity of the organisms and by the
various forms of biological interaction in communi-
ties.
The numerous types of interaction within a com-
munity and their physical environment mean that
there is positive and negative feedback in (com-
munity) ecosystems, which can have a different
specific form in each community. However, self-regu-
lation as a sort of steering force does not exist; this is
an idea which arises from an organismic view of
ecosystems. The dilemma of holistic ecosystems re-
search is clearly demonstrated in cities, since most
urban ecosystems, in particular the 'ecosystem city',
are not self-regulating, but are 'regulated' by humans.
Some people reject the ecosystem concept completely
because it is so closely associated with the holistic
approach to ecosystem research.
In terms of Tansley's original definition it is poss-
ible in the case of cities to define both individual
urban 'ecosystems' (e.g. the centre, parks, wasteland)
as well as 'the ecosystem' of the entire city (e.g.
Brussels). The important thing is not the mere use of
the term ecosystem, but the way in which it is used.
The 'ecosystem' of a specific city is in fact a complex,
consisting of various communities which can overlap
and interact to a greater or lesser extent. A city park
itself can be divided into various sub-communities
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Urban ecology 175
(e.g. lawns, meadows, woodlands, streams, ponds),
which can in turn interact.
The variety of urban activities and land uses means
that the boundaries between various sub-habitats are
particularly marked, but individual populations and
communities can stretch over several of these sub-
habitats. The boundaries of a survey area ('an ecosys-
tem') are laid down by the scientist more or less
arbitrarily, depending on the topic under study and on
practical considerations. Thus the investigation of the
energy balance of an entire city will make use of the
politically defined city boundaries, even if this en-
closes non-urban settlements. In other cases, such as
the comparison of species distribution between built-
up and rural areas, other boundaries are ecologically
more appropriate.
MIGRATION, DISPERSAL AND EXTINCTION OF
SPECIES
A significant and far-reaching aspect of global ecolog-
ical change is the elimination of biogeographical bar-
riers to the dispersal of species as a result of human
activities (Elton, 1958; Drake et al., 1989; D'Antonio
& Vitousek, 1992). The result of this is that species
are migrating widely between continents and islands,
meeting species they would otherwise never have
encountered. This applies in particular to habitats
which have been considerably altered, or indeed cre-
ated by human activity. Thus it is much more com-
mon in cities than in natural habitats to find
communities consisting of species without a common
evolutionary past which have not previously been
found elsewhere in such a combination.
A relatively small proportion of the species inten-
tionally or unintentionally introduced by humans from
other geographic regions successfully disperse and
establish themselves in relatively natural communities
(see Lohmeyer & Sukopp (1992) for agriophytes in
Central Europe, D'Antonio & Vitousek (1992) for
grasses globally).
In addition to overcoming geographical barriers,
anthropochorous dispersal can also overcome ecologi-
cal barriers when species meet under new conditions.
Thus Urbanska (1992) reports about the two thistles
Carduus acanthoides L. and Carduus nutans L. in
Ontario, which differ in their ecological demands. C.
acanthoides prefers in Ontario skelletal, permeable
soils, whereas C. nutans prefers moister sites. Anthro-
pogeneous dispersal meant that both species, which
had been separated by ecological preferences, met on
a site created by human activity - a roadside embank-
ment - where they crossed. By back-crossing, the tol-
erance range of C. acanthoides was considerably
increased.
In the literature on introduced species and their
distribution, the terms 'invasion' and 'colonization'
are often used synonymously (e.g. Ehrlich, 1986,
1989). With regards to both the anthropochorous spe-
cies from other biogeographic regions, and also to the
process of colonization in primary successions (and in
part in secondary successions) it is, however, sensible
to distinguish between the terms, or at least to differ-
entiate between the processes covered by the term
'biological invasion'.
For plants, Bazzaz (1986) distinguishes between
.colonizers', 'immigrants', and 'invaders'. Colonizers
settle open sites in primary succession, and in part in
secondary succession. A common feature of all colo-
nizers is that the site they settle is not yet occupied by
other species, and that there is initially no compe-
tition. If there are repeated disturbances it may not
come to competition between species at all. This is a
common situation in urban and industrial areas.
'Immigrants' are species which do not significantly
impede populations already to be found on a site, and
which integrate themselves into the existing com-
munity. 'Invaders' are species which penetrate natu-
ral, intact phytocoenoses, and either dominate these or
displace other species. Such situations are very im-
probable in urban habitats since these are either
influenced or created by human activity.
The characteristics described by Bazzaz (1986) for
immigrants and invaders corresponded more or less to
the processes of 'integration' and 'displacement', or
the behaviour of agriophytes in 'unsaturated' and
,saturated' plant communities as described by Sukopp
(1962). It is, however, appropriate to examine sepa-
rately the process described as colonization, and to
separate this clearly from the processes of 'inte-
gration' and 'displacement'.
Of the many species introduced by human activity
from other biogeographic regions, only some are suc-
cessful (see Lohmeyer & Sukopp, 1992 for ferns and
flowering plants in Central Europe, Simberloff, 1986
for insects). According to Lohmeyer & Sukopp
(1992), in near-natural vegetation in Central Europe
less than 2% of the introduced plant species have
established themselves. (On the question of 'Resist-
ance of plant communities to biological invasion' see
also Trepl, 1990, 1994; Trepl & Sukopp, 1994.)
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176 Franz Rebele
In cities, the proportion of successfully established
introduced species in the flora is higher than in near-
natural habitats (Kowarik, 1991). These belong
mainly to the group of colonizers, or are integrated in
plant communities which have been newly formed
under urban conditions (many ruderal plant communi-
ties).
Not only the introduction and dispersal of species
which were not previously found in an area (including
not only 'alien' species but all synanthropic species),
but also the local extinction of species is typical for
towns and cities. In particular this affects species from
communities on natural or near-natural sites, i.e. habi-
tats which are classically non-urban.
The reasons for this are closely linked to urban
development. Construction work directly destroys
habitats, eliminating local populations (see Rebele,
1991). Other processes, such as the lowering of
ground water levels or eutrophication, can change site
factors and thus lead to changes in the abundances of
populations, even to the extent of elimination (see
Sukopp, 198 1).
However, where there are Ino alterations of the
habitat, direct displacement by invasive competitors
leading to local extermination of species is very un-
likely in cities. Firstly, in cities there are a large
number of habitats in which resources are not limited,
or in which the low population density means that
there is no competition for limited resources. This is
the case for a number of plant species which settle
open habitats (colonizers, see above), but also for the
large group of phytophagous insects. On the other
hand, a precondition for the exclusion of competition
is stable conditions, which are rarely, if ever, achieved
in urban habitats (see also stability and equilibrium, p.
182).
According to MacArthur & Wilson (1967), islands
have a natural turnover of species. The number of
species on islands is determined by an equilibrium
between immigration and extinction. For cities or
individual urban communities the turnover of species
ought to be particularly high at least for certain groups
of organisms, since the conditions exist for both im-
migration (in the wider sense) and local extermi-
nation. The processes underlying the turnover of
species, however, are not determined primarily by the
ability of the species to disperse and by biological
interactions, but by introduction, dispersal and local
extermination by human activity. In contrast to the
assumption of stability in the island model of
MacArthur & Wilson (1967), there is not generally a
balance of species richness in cities. The numbers of
species can either increase or decrease.
In Zurich the number of flowering plant species
increased from 970 to 1200 species over the last 150
years. One hundred and thirty species died out, but
about 350 introduced species became established
(Landolt, 1992). A decline in the absolute number of
species may occur, if besides the process of extermi-
nation of species in near-natural habitats habitat diver-
sity there is diminished through enforced sealing of
wasteland or levelling of sites (Rebele, 1991). For
Bochum (Ruhrgebiet) a decline of the flora of vascu-
lar plants from 600 species in 1887 (Hamann, 1976)
to about 580 in 1985 (Schulte, 1985) can be recog-
nized.
HABITATS, ENVIRONMENTAL FACTORS AND
RESOURCES
In urban and industrial areas it is possible not only to
find many 'exotic' species, but also substrates and
habitats which are alien to the original states. The
introduced substrates can be both natural substrates
from other geographical and geological regions (e.g.
railway ballast, mine spoil), or artificial substrates
(e.g. slag), or a mixture of artificial and natural sub-
strates (e.g. slag/clay). In extreme cases earth can be
transported even between continents (Lindroth, 1957).
A large number of urban habitats such as buildings,
roofs, underground pipes are specifically related to
human habitation. Natural soils are usually consider-
ably altered by human settlement activities and pro-
duction (soil is moved, substrates are mixed, thre is
compaction and sealing over). An overview of the
properties and the classification of soils in cities is
given by Hollis (1991). Various technological sub-
strates of urban and industrial sites are listed by
Meuser (1993).
A feature of many cities is the wide variety of
environmental conditions. Soils in cities can be very
poor in nutrients, or may be highly enriched (e.g.
former sewage farms). Toxicity can play a role in
places, due to deposits of toxic substances and sub-
strates, or as a result of pollution. There are open sites
with high relative irradiance and very dark habitats
(artificial caves). Habitats can also be anything from
very dry to wet, although dry sites are more common
in urban habitats. For plants the relationship between
light and soil resources is of vital importance. Sub-
strates with low levels of available nutrients tend to
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Urban ecology 177
offer ample light, whereas if soil resources are plenti-
ful there is usually a lack of light at ground level
(Tilman, 1986).
The urban climate is generally warmer than that in
the surrounding countryside. Even more significant
for organisms than the annual mean temperature,
however, are microclimate variations. Temperatures,
for example, can vary extremely widely over short
distances. Abiotic site factors can be relatively homo-
geneous over areas (e.g. large excavations or embank-
ments), but also very heterogeneous (e.g. industrial
wasteland).
Extreme abiotic site conditions (e.g. lack of light or
water) are an important selection factor for the estab-
lishment of many plant and animal species. Begon et
al. (1990), however, rightly point out that 'extreme'
sites are only 'extreme' to the outside observer. Lim-
ited resources can be a stress factor for some species,
leading to reduced productivity. For tolerant species,
on the other hand, the conditions can be near ideal,
since they do not have to face the competition of other
species for resources.
DISTURBANCE
According to White & Pickett (1985), a disturbance is
a relatively discrete event which suddenly disrupts the
structure of an ecosystem, community or population,
changing either the availability of resources or the
physical environment. This is a very broad definition,
but it says nothing about the cause or the nature of the
disturbance.
Disturbances, can have not only physical but also
biological causes and generally lead to the death or
reduction in abundance of a species, which can in turn
favour other species. Thus Sousa (1984) speaks of
'discrete, punctuated killing, displacement, or damag-
ing of one or more individuals (or colonies) that
directly or indirectly creates an opportunity for new
individuals (or colonies) to become established' (p.
356).
However, disturbances in ecosystems do not
necessarily lead to the death of organisms. Soil distur-
bances, for example, can improve the conditions for
germination, or increase the availability of nutrients.
Local eutrophication can also represent a sudden alter-
ation in the availability of resources, although not if
this is a continual state. As a rule, however, both
physical and biological disturbances involve a de-
struction of biomass, the physical generally being
non-specific, but the biological often being specific,
namely in the form of predation.
Natural disturbances include storms, fires and land-
slides, but also the impact of grazing animals. In
towns and cities there are a number of additional
disturbances, particularly those linked with construc-
tion activities, gardening and recreation. Anthropoge-
neous disturbances are usually of much greater
importance in urban habitats than natural disturbance
factors.
Disturbed areas in cities can vary in size from a few
square centimetres to square kilometres. It is particu-
larly common for construction work to destroy com-
munities completely. Other disturbances are localized,
and less severe. The extent and the intensity of distur-
bances has an important influence on its biological
effectiveness. Localized disturbances can increase the
heterogeneity of the environment, creating a larger
number of safe sites for plant species (Harper, 1977;
Grubb, 1977). More wide-spread disturbances, such as
the removal of top soil prior to construction, can lead
to a reduction in the heterogeneity. Frequently recur-
ring disturbances can hinder the growth of perennial
plant species, and thus also reduce heterogeneity.
The regional frequency of anthropogenous distur-
bances depends primarily on economic, political and
social factors, which means that in cities they are very
difficult to predict, since the economy does not de-
velop linearly, social habits and fashions change con-
stantly, and political processes are often chaotic.
Various statements have been made, for example,
about the lifetime of buildings. 'Turnover rates', how-
ever, cannot be generalized for all cities, and can even
vary considerably within a city, as well as depending
on economic and political developments. It is gener-
ally true for urban ecosystems that disturbances do not
usually occur in phases, but displaced, and thus vari-
ous stages of succession exist side-by-side. An im-
portant feature of species in urban habitats is that they
are able either to regenerate themselves after distur-
bances or to settle in newly created gaps.
Here, typical urban-industrial ecosystems are con-
sidered. Special consideration is given to the numer-
ous disturbances to residual near-natural sites within
and directly around cities.
COLONIZATION OF OPEN HABITATS
Previously, the process of colonization was con-
sidered in connection with introduced plant species.
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178 Franz Rebele
However, some of the most abundant and successful
colonizers are native species, though these have usu-
ally developed their characteristics under different
evolutionary circumstances. As a rule these are spe-
cies from naturally disturbed habitats, e.g. river banks.
In the process of colonization, newly created habi-
tats are settled, or sites are re-settled following a
disturbance. It is not usual in the case of a secondary
succession to distinguish between a settlement by
migrant diaspores from elsewhere and the recruitment
of plant populations from the diaspore bank. Thus for
Grubb (1987), for example, colonizers are simply 'the
first organisms to become established in a succession'
(p. 81).
The new colonization of open sites is primarily
dependent on the number of diaspores reaching a site
or already to be found there, and on the physical-
chemical environmental factors. For example, plant
populations must be able to germinate and establish
themselves. Competition within and between species
does not occur as long as relevant population densities
are not exceeded. Other forms of biological interac-
tion can play a role, such as predation by seed-eating
birds or insects. In addition to the establishment of
new organisms from seed etc., some plant species can
also settle sites by vegetative spread.
In cities, open habitats (those not already occupied
by other species) are relatively common. This is due
to the creation of new habitats and the disturbances
already considered in the previous two sections. The
colonizers of open sites use up the existing soil nutri-
ents and thus create zones of resource depletion. On
the other hand they also produce organic debris which
returns nutrients to the soil. In this way they produce
a spatial heterogeneity of plant nutrients even in rela-
tively homogeneous substrates.
Animals (including e.g. ants or earth wasps) can
also contribute significantly to the restructuring of the
micro-habitats of open sites such as sand heaps.
SUCCESSIONS
Successions are directed changes of the species com-
position over time. Vegetation successions are gener-
ally also linked to a clear change of the vegetation
structure. A distinction is usually made between pri-
mary and secondary successions.
Primary successions occur naturally on sand dunes,
glacial debris or volcanic ash. Due to man's industrial
activities, primary successions can also occur in cities.
They occur wherever natural soil is excavated or a
substrate is spread which did not previously bear any
plant growth and thus is devoid of any diaspore bank.
The content of organic substances and nitrogen is
initially very low in a primary succession. Other
nutrients are not limited in many substrates (Tilman,
1986). In the course of primary succession, organic
matter and nitrogen accumulates. The level of avail-
able phosphates, on the other hand, can decline (Vi-
tousek & Walker, 1987).
There is usually an increase in the number of
species in the first years of primary vegetation succes-
sion, which then levels off in the course of succession
(Crawley, 1986). In cities the settlement of primary
habitats by plants is often more rapid than in isolated
natural primary successions, since there are usually
large numbers of diaspores of colonizing species in
the near vicinity. Birds perching in adjacent gardens,
parks, road sides or wasteland can also accelerate the
colonization process (van der Pijl, 1982; McDonnell
& Stiles, 1983).
Secondary successions occur when sites which al-
ready bore vegetation cover are disturbed (storm dam-
age in forests), or when cultivated land is left to lie
fallow. In cities, secondary successions are particular
frequent, since both disturbances and also temporary
fallow phases are very common. Features of second-
ary successions are the presence of organic matter at
the start and an increased release of nutrient after a
disturbance of the vegetation cover (or direct enrich-
ment with dead organic matter).
The vegetation development on deposited top-soils
and landfill-soils in cities usually has the character of
a secondary succession, or a mixed-form of secondary
and primary successions. This is because substrates
are often used which have already had vegetation
cover (soil recycling), or which are enriched with
organic matter and diaspores (e.g. compost enriched
top-soils).
A further important feature of secondary succes-
sions is the 'initial floristic composition factor' (Egler,
1954). Most species which play a part in the course of
succession (annuals, short- and long-lived perennials),
are already present at the start of the succession, or
migrate to the site from adjacent areas soon after the
disturbance or the deposition of the substrate. De-
pending on the nutrient content of the soil, the veg-
etation develops faster or slower, or one can recognize
successional stages (dominance of annuals, mono-
carpic perennials, polycarpic perennials). Secondary
succession on urban deposit soils usually have the
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Urban ecology 179
greatest number of species at the beginning of the
succession. The higher the level of nitrogen in the
applied substrate the more rapid is the decline in the
species density, since in highly productive plant com-
munities the lack of light at the soil surface hinders
both the early successional species as well as the
woody perennials (Rebele, 1992).
In urban and industrial areas, the frequency of
disturbances means that early- and mid-successional
stages are common. Late successional stages, on the
other hand, are very rare (e.g. birch forests on mine
spoil heaps in the Ruhrgebiet, or some 'urban forests'
on ruderal sites in Berlin).
BIOLOGICAL INTERACTION
Most communities are organized, according to Sousa
(1984), by biological interactions such as competition
and predation in the wider sense of the term, as well
as by disturbances (see also Begon et al., 1990).
Mutualism can also be important. The relative import-
ance of the various forms of biological interaction
can, however, undergo a systematic change, with
competition and predation being less important in
mechanically disturbed habitats. Disturbances have
already been considered, see page 177. Some forms of
biological interaction are considered here in more
detail.
(a) Competition
There are several different kinds of competition. Re-
source or exploitative competition, which is particu-
larly important for plants, requires niche
differentiation if it is not to lead to competitive ex-
clusion. Interference competition and competition for
space, on the other hand, do not require niche differ-
entiation (Yodzis, 1986).
Exploitative competition occurs at the same trophic
level and the same resources demands. According to
the classical competition theory, two species with
exactly the same resource requirements cannot co-ex-
ist (Gause, 1934). The co-existence of species at the
same trophic levels can be explained by niche differ-
entiation, which is either produced by current compe-
tition, or already exists because of evolutionary
avoidance of competition in the past ('ghost of com-
petition past', Connell, 1980). A feature of co-existing
species in the case of niche differentiation is that the
intra-specific competition is more important for the
dominant species than the inter-specific competition.
Some urban habitats demonstrate considerable het-
erogeneity over time and space. For autotrophic
plants, light and nitrogen, for example, are the two
crucial factors which can be at a minimum and which
are the subject of competition. Even within relatively
homogeneous old fields the nitrogen content in an
area of 0.14 ha can vary be a factor of 3-6.7 between
samples. During secondary succession, the nitrogen
content can also vary by a factor of 6 within 120 years
(Tilman, 1986).
The species which grow together on fallow land
need not form part of a converging community. Each
species can successfully compete with other species
along a nutrient/light gradient (Tilman, 1986). Even
when there is exploitative competition, the spatial and
temporal environment heterogeneity can ensure the
coexistence of competing species.
Niche differentiation in the narrow sense of the
term is, however, more of an ideal case, and requires
a relatively stable environment and a long process of
joint evolution, or evolution under similar environ-
mental conditions (Crawley, 1987). Even under natu-
ral conditions, without human influence, this is not
always so, or is controversial (see Hubbell & Foster,
1986). Under urban conditions it is even less the case,
since unforeseeable disturbances make the environ-
ment extremely unstable.
In the case of competition which requires no niche
differentiation or environmental heterogeneity, the
initial density of a species can be of significance, or
first occupancy of the site (founder effect, see e.g.
Schmidt, 1981; Rebele, 1994). Since gaps are repeat-
edly being created in urban communities by physical
disturbances (in part also by predation), there are
always competition-free situations which allow
weaker species to co-exist with superior competitors.
(b) Predation
Of the various forms of predation, herbivory is con-
sidered here.
There are numerous examples of 'empty niches' for
herbivores in urban habitats in particular. Competition
between herbivores generally plays a less important
role, particularly in the case of phytophagous insects
(Hairston, Smith & Slobodkin, 1960; Slobodkin,
Smith & Hairston, 1967; Strong, Lawton & South-
wood, 1984; Jermy, 1985). Investigations of herbivo-
rous insects on bracken (Pteridium aquilinum (L.)
Kuhn) on three continents have shown that some
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180 Franz Rebele
feeding sites of this wide-spread, cosmopolitan spe-
cies are exploited in one part of the world but not in
another, since the insect fauna differs (Lawton, 1984).
Comparisons are frequently made between the phy-
tophagous fauna of native and exotic species. It is
known that neophytic species in their anthropochoric
areas usually have a lower number of herbivorous
insects than in their area of origin (Lohmeyer &
Sukopp, 1992). It has also been established for entire
plant groups, such as woody perennials, that exotic
species represent a poorer source of nutrition for
animals than native species (Southwood, 1961;
Kennedy & Southwood, 1984; Kowarik, 1986). How-
ever, the difference is probably not as large as is
generally supposed (see, e.g. Glauche, 1991). Thus
Owen (1991), after observing her garden in Leicester
for 15 years, concluded that only 33.7% of all native
plant species served as a source of food for moth
species, compared with 42.4% of all exotic species.
However, of a total of sixty-eight moth species, forty-
six fed on native species compared with thirty-eight
on exotic species.
There are a number of possible reasons for the
difference in the phytophagous fauna, and urban habi-
tats offer ample opportunity to investigate these. The
feeding on exotic plant species by phytophagous in-
sects can depend, for example, on the length of their
presence in the new area, the distance to the area of
autochtonous distribution, the frequency of the species
in their area of origin as well as in their anthropoge-
nous area, the total size of the area, the taxonomic
proximity of the species to native species, or the
taxonomic status of the species (families with numer-
ous species also have a relatively large number of
phytophages). One of the crucial factors is probably
the existence of transport and trade routes, and the
frequency and volume of the exchange of goods (see
e.g. Simberloff, 1989).
It is certainly misleading to compare native plant
species with rich phytophagous insect fauna with ex-
otic species which are poor in phytophagous fauna.
There are also large differences between native spe-
cies as far as the numbers of phytophagous insects is
concerned (e.g. Quercus species/Taxus baccata L. in
Central Europe). Some introduced species have low
numbers of phytophagous insects in their place of
origin also (e.g. Ailanthus altissima (Mill.) Swingle,
after Rohricht, 1991). As the example of Pteridium
aquilinium shows, the use of plant species can also
vary widely over its area of natural distribution.
Of the other herbivores, common urban species are
of particular interest, such as rabbits in many central
European cities. Herbivores cause biological 'distur-
bances' (see above) amongst the plant species, and
can thus affect biological interactions such as compe-
tition. It is known that rabbits can have a considerable
influence on the species diversity of chalk grassland
(Tansley & Adamson, 1925).
Many carnivorous species change their feeding
habits in towns and cities. A well-known example is
for fox populations in Britain (Harris, 1986). But
population densities and life expectancy can also be
affected, and with them the age structure of popula-
tions (Harris, 1986).
(c) Mutualism
The majority of the earth's biomass consists of organ-
isms which have mutualist relationships to others
(Begon et al., 1990). Well-known examples of mutu-
alism are the symbiosis of legumes with nitrogen-
fixing bacteria. Legumes also play an important role
in urban phytocoenoses, e.g. on N-poor sandy soils.
They are not always among the first colonizers, but
they are among the species which form the dense
ground cover.
In natural ecosystems, mycorrhiza is the rule rather
than the exception. In some families, e.g. Brassi-
caceae, Carvophvllacea and Chenopodiaceae, mycor-
rhiza is rarer, but in the case of woody species,
grasses and ferns it is widespread. The VA-mycor-
rhiza common in some families serves mainly the
uptake of phosphates (Harley & Harley, 1987). Phos-
phates often act as a constraint for later successional
stages (see above).
Annuals are frequently without mycorrhiza. This is
true both for annuals on ruderal sites rich in nitrogen,
as well as for therophytes on dry sandy lawns and
spoil heaps (e.g. Cerastium semidecandrum L., Chae-
narrhinum minus (L.) Lange, Minuartia verna (L.)
Hiern (see Harley & Harley, 1987). It is still question-
able whether mycorrhiza plays a role right from the
start in the colonization of urban habitats by grasses
and woody perennials.
Where soil is removed and stored, the microbial
activity and the proportion of VA-mycorrhiza declines
and this also delays the development of mycorrhiza in
grasses, or reduces its level (Rimmer, 1991). Reeves
et al. (1979) report that the level of mycorrhiza on
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Urban ecology 181
severely disturbed sites is lower than on natural,
undisturbed sites.
Overall, it seems that symbiosis with nitrogen-
fixing bacteria plays an important role for urban habi-
tats in view of the generally low levels of nitrogen,
whereas mycorrhiza plays a less important role than,
for example, in forests.
(d) Interactions between interactions
Thompson (1988) has pointed out that there are no
sharp distinctions between the various forms of bio-
logical interaction, that competition and mutualistic
interactions overlap. Thus, if competing plants blos-
som at the same time they will attract more pollina-
tors, to the advantage of both species. The symbiosis
between legumes and N-fixing bacteria is only of
advantage for legumes on N-poor soils, but not on
N-rich soils. The results of interaction can also be
dependent on environmental factors. Several refer-
ences have already been made to predation as a bio-
logical 'disturbance' which reduces competition.
SPECIES RICHNESS AND DIVERSITY
Species richness can be taken to mean the number of
species per unit area (species density), or in a specific
location (numerical species richness). When calculat-
ing diversity, the evenness plays a role, and it can be
expressed using various indices (Hurlbert, 1971; Peet,
1974). After Whittaker (1965), the diversity can be
divided into several components. Alpha-diversity
refers to the diversity within communities, beta-diver-
sity to the rate of change in the species cornposition
along a habitat gradient (diversity between communi-
ties), and the gamma-diversity to the overall diversity
of the landscape (see also Peet, Glenn-Lewin &
Walker Wolf, 1983).
The alpha-diversity in urban phytocoenoses can
vary from very low to very high. Examples of plant
communities with low diversity can be found in ex-
treme soil conditions such as very low or very high
pH, on toxic substrates or also under eutrophic condi-
tions (e.g. dominant stands of Calamagrostis epigejos
(L.) Roth or Urtica dioica L. Examples of high alpha-
diversity are to be found in some ruderal plant com-
munities with a species density of more than 40
species per square metre with a low dominance of
individual species. If one compares various phyto-
coenoses in urban habitats, then it is not rare to find
plant communities with very low diversity as close
neighbours of communities with high diversity.
The beta-diversity can be very high in cities, since
very pronounced gradients are frequently formed
there. There are, however, also some very uniform
areas. The fact that cities are frequently richer in
species than the surrounding areas (at least as far as
plant species and some groups of animals are con-
cerned, see Klausnitzer, 1993; Landolt, 1992; Reich-
holf, 1989; Sukopp & Trepl, 1993, Wittig, 1991) can
be attributed above all to the great habitat diversity.
The relationship between introduced species which
are newly establishing themselves to species facing
local extermination also plays a role.
With reference to both the habitat diversity and also
the species turnover in cities, there are not only gen-
eral characteristics, but also geographical and histori-
cal components. Scandinavian cities differ from
central European cities, and the latter from Mediter-
ranean cities, not only in their architecture, but also in
the species composition. Within central Europe, cities
of comparable size can also differ in their species
abundance, with historical components (urban devel-
opment, trade connections, industrial development
etc.) providing a degree of 'local character' (Gilbert,
1989).
In addition to (spatial) habitat diversity, the time
factor also plays an important role, with various suc-
cessional stages side-by--side. References were made
to vegetation succession earlier (p. 178). Little is
known as yet about succession of soil communities
and animal communities in urban ecosystems.
For some animal groups it is not the species rich-
ness of plant communities which is most important,
but the structural diversity of vegetation, and/or the
spatial heterogeneity of the non-living environment
(e.g. for birds or lizards).
It should be noted that the floral diversity can either
increase or decrease in the course of successions,
depending on the type of successions. Regarding soil
resources, the highest diversity is generally found on
sites of moderate fertility (Tilman, 1982). The size
and nature of disturbances also have an influence on
species diversity, with the highest levels frequently
being found where disturbances are intermediate, an
observation not only made in towns and cities ('inter-
mediate disturbance hypothesis', Connell, 1978). Ob-
jections have, however, been voiced against making
generalizations in this context (Yodzis, 1986; Leigh,
1 990).
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182 Franz Rebele
COMPLEXITY
Tansley (1935) pointed out that the degree of inte-
gration of ecosystems can vary. This depends in the
first place on the nature and extent of interactions
within the community. According to Pimm (1984) a
community has the following components.
1. The species richness.
2. The connectance (the number of inter-specific
interactions as a fraction of all possible inter-
specific interactions).
3. The interaction strength (i.e. the effect of the
density of a species on the growth rate of an-
other species).
4. The evenness within a community.
Little work has as yet been done on the complexity
of urban communities. However, it is not possible to
set up a simple rule, for example that urban communi-
ties are less complex than non-urban ones. Rather, the
few investigations in the literature suggest that there is
a whole range from very low to high complexity
(Briand, 1983, example of aspen parkland), as does
the rich fauna and flora of much wasteland.
On average, the complexity of urban communities
is probably less than various non-urban ecosystems
(e.g. forests), since biological interactions, especially
inter-specific competition becomes less significant
given the frequent disturbances.
STABILITY AND EQUILIBRIUM
Tansley (1935) also pointed out that the ecosystems
are less stable than, for example, chemical elements,
since the ecosystems consist of components which are
themselves unstable to a lesser or greater extent - the
climate, the soil, and the organisms. The climate
changes with time, soils develop, and the composition
of organisms can also change, even in natural ecosys-
tems undisturbed by humans. Since, by definition,
ecosystems are not enclosed systems, components
from other systems can penetrate them and cause
them to disintegrate. Under natural conditions a dy-
namic equilibrium may be established over decades or
centuries in the sequence of successions, but there are
also forces which work against this.
A major problem in the discussion of stability is
that this was originally defined in terms of natural
ecosystems in equilibrium. According to Hurd et al.
(1971), stability is a concept adopted fom thermody-
namics, and defined as 'the ability of a system to
maintain or return to its ground state after external
perturbation' (p. 1134). The definition of Holling
(1973) is similar, but using 'equilibrium state' and
'temporary disturbance', and omitting the aspect of
maintaining its state: 'Stability, ..., is the ability of a
system to return to an equilibrium state after a tempo-
rary disturbance. The more rapidly it returns, and with
the least fluctuation, the more stable it is.' (p. 17)
Pimm (1984), corresponding essentially with May
(1973), defines stability more precisely: 'A system is
deemed stable if and only if the variables all return to
the initial equilibrium following their being perturbed
from it. A system is locally stable if this return is
known to apply only certainly for small perturbations
and globally stable if the system returns from all
possible perturbations'. 'Resilience', for Pimm (1984)
expresses, 'how fast the variables return towards their
equilibrium following a perturbation', and is therefore
defined only for a stable system, but not for unstable
ones. 'Resistance', on the other hand, is 'the degree to
which a variable is changed following a perturbation'
(p. 322).
The variables which play a role in ecosystems are,
for example, the number of species, the abundance of
the various species, the productivity, element concen-
trations or energy levels. Common to the definitions
presented here is the assumption of a ground state or
equilibrium. For living communities, an equilibrium
means that the net growth rate of populations (but also
other parameters such as biomass and element con-
centration) are zero.
Because of disturbances and climate fluctuations,
natural communities are seldom in a state of equilib-
rium, except perhaps if viewed on a wider spatial
level. Chesson & Case (1986) point out that many
inequilibrium theories are compatible with the theory
of global stability of a community. Local inequilibria
can lead, over the average of all local communities to
a global stable equilibrium. This also corresponds to
the concept of 'shifting mosaic steady-state' presented
for natural forest ecosystems by Bormann & Likens
(1979).
When considered over larger areas, however, natu-
ral systems are also frequently in a state of transition
(depending on the extent and the intensity of the
disturbance). Connell (1978) gives examples of natu-
ral ecosystems (tropical rainforests, coral reefs),
which are in states of transition after a disturbance.
Turning to urban ecosystems, this means that states
of equilibrium are virtually impossible, since it is
highly probable that a perturbed system will be dis-
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Urban ecology 183
turbed again before the state of equilibrium has been
reached. It will also rarely be possible to speak of a
system returning to equilibrium, since the systems
disturbed in a town or city will rarely be in a state of
equilibrium to start with, but in a state of transition or
a state maintained by human intervention (such as
regular mowing and fertilizing of lawns).
It follows from this that when assessing populations
in urban communities, and biological interactions be-
tween them, theories and models which are based on
equilibria and stable environments will be less rel-
evant than for systems with relatively constant envi-
ronments and little disturbance.
A definition of stability which also includes states
maintained by humans is given by Gigon (1984):
'Ecological stability is the persistence of an ecological
system and its ability to return after alteration to its
initial position' (p. 14).
In this definition, the term 'equilibrium' is replaced
by 'initial position', so that the return to a state
maintained by human activity can be termed 'stabil-
ity'. According to Gigon (1984), if a meadow was no
longer mown then 'instability' would result. However,
the succession from a fallow meadow to a forest can
be regarded as the tendency towards a natural point of
equilibrium. It is therefore essential when discussing
stability, to explain what one means by the stability of
an ecological system, or which stable state is envis-
aged (e.g. the meadow or the 'natural equilibrium').
The question of equilibrium is usually dealt with in
conjunction with the complexity and diversity of com-
munities. As has been noted, high diversity is found
more frequently in instable communities (Connell,
1978), and complex communities are not necessarily
more stable than less complex ones.
PRODUCTIVITY
The productivity of urban ecosystems can vary from
very low to very high depending on soil parameters.
Low productivity is found, e.g. in plant communities
on slag heaps, gravel, or other dry substrates poor in
nutrients. Very high productivity can be found in early
to mid-successional stages on eutrophic soil deposits.
In Brussels, ruderal phytocoenoses such as Tanaceto-
Artemisietum vulgaris or Solidaginetum giganteae
achieve a yearly productivity of 14-20 tonnes/hectare
(Duvigneaud, 1975).
If a city is examined as a whole, then the productiv-
ity depends primarily on the degree of land sealing, or
put another way, on the proportion of area which can
be settled by green plants. In some cities the unsealed
area can be as much as 50% of the total surface area,
e.g. Brussels (Duvigneaud & Denayer-de Smet, 1977)
or Halle-Neustadt (Frotscher, 1990). Within the cities
there are usually different zones for the level of
sealing. The city centre is usually more sealed than
the outer districts (see e.g. B6cker, 1985).
The standing biomass is not only dependent on the
proportion of the area available to plants for settle-
ments, but also on the age of the successional stage.
The older the successional stage, then the more
biomass is available. Little data has been published
about the overall phytomass in cities. The figure of
750,000 tonnes dry matter given for Brussels (Duvi-
gneaud & Denayer-de Smet, 1977) corresponds over
the entire area of 162 km2 approximately to a phy-
tomass of 46 tonnes/hectare.
DECOMPOSERS AND DETRITIVORES
Decomposers and detritivores are very important for
the formation of soils, and they play an even more
important role in cities. A special feature of urban
ecosystems is that in addition to the production of
dead organic matter by the urban organism communi-
ties, organic waste is also produced by humans (usu-
ally imported from outside). To some extent this is
removed again by waste disposal, emissions, sewage
systems and run-off into running waters, but there is
also an accumulation of nutrients in certain places in
the cities from antropogenic sources.
CONCLUSIONS
The topics discussed in this paper are questions of
general importance in ecology. Nevertheless, there are
some special features of urban ecosystems like mosaic
phenomena, specific disturbance regimes, the pro-
cesses of species invasions and extinctions, which
influence the structure and dynamics of plant and
animal populations, the organization and characteris-
tics of biotic communities and the landscape pattern
as well in a different manner compared with natural
ecosystems. On behalf of the ongoing urbanization
process, urban ecosystems should attract increasing
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184 Franz Rebele
attention by ecologists, not only to solve practical
problems, but also to use the opportunity for the study
of fundamental questions in ecology.
ACKNOWLEDGMENTS
I would like to thank Prof. Dr Ludwig Trepl (Munich)
for his valuable comments on the manuscript, and
Richard Holmes for help with the English.
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