bram mabelis
Has Lasius neglectus invaded the Netherlands ? |
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Occurance Lasius neglectus in Europe |
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Introduction During the last decades the built-up area of Warsaw expanded very fast. As a result several green areas disappeared, while the remaining ones are getting more and more under urban pressure. Consequently, more green areas will disappear (Szulczewska and Kaliszuk 2003). This will affect the liveability of the city. Green areas are important for improving the urban climate: trees and bushes increase air humidity, give some cooling on hot days, produce oxygen and filter the air. Moreover, urban green retains rain-water and its soil has important cleaning properties: some noxious substances can be broken down, at least if the soil is not too much polluted. Urban green has not only an ecological function, but is also important for recreation. For both functions it is important to keep biodiversity at a high level. The Convention of Biological Diversity (Rio de Janeiro 1992), which was signed by many countries, aims at the maintenance of ecosystem, species and genetic diversity. This Treaty also has consequences for local communities, according to Local Agenda 21. The city council can meet its obligations by establishing a sustainable green infrastructure. The maintenance of such a robust green infrastructure is not only important for the survival of native plants and animal species, but also for maintaining a healthy environment and for giving citizens the opportunity to relax and experience nature. In order to prevent alienation of citizens from nature the presence of native species should be favoured above introduced exotic species, although the aim to maintain a high level of biodiversity in a city emanates mainly from the wish to keep it liveable and healthy (Kelcey 1978, Trojan 1981a, Goode 1989). Ecological qualities of urban nature
should be guaranteed despite urban processes. Therefore, a strategy should
be followed in which water, energy and waste flows are managed properly
(Tjallingii 1993, 1995, 1996). Moreover, fragmentation of urban green should
be prevented as much as possible. Generally, urban green is less fragmented
in the outskirts of the city than in the centre, where green areas are
generally smaller and more isolated by roads and buildings.
Urban environment In flat land a city like Warsaw rises up as a rock island. Buildings with holes, crevices and hiding places can be suitable for rock-dwelling birds and bats to hide, reproduce or hibernate, while old walls (with calcium- rich cement between the stones) can be suitable as habitat for wall plants. Moreover, cellars, which are constant cool and moist, add a kind of cave environment to the city. Several species use cellars to hibernate, like some beetles and night butterflies. A city can also be considered as
a heat island: the average temperature is higher in a city than in its
surroundings, despite the fact that the sun will shine here less often
and less fierce than in the countryside, due to air pollution. However,
a great deal of the city surface is covered by buildings, asphalt and flat
stones, which can preserve heat for a long time. The relative high temperature
of the city is suitable for thermophilic species. Consequently, the northern
limit of the distribution area of several thermophilic species is situated
in cities. Examples are known of plants (Sukopp and Wurzel 1995), digger
wasps (Whiteley 1994), longhorned beetles (Burakowski and Nowakowski 1981b),
rove beetles and glow worms (Klausnitzer 1989) and springtails (Sterzy?ska
1982), among others. Thanks to the warmer city climate birds can produce
more clutches of eggs per year (Klausnitzer 1989). Moreover, winters are
less severe in cities. Consequently water will freeze less easily, which
may be profitable for water birds. On the other hand, the warmer climate
of a city is disadvantageous for some insect species: e.g. a parasite can
miss the connection with its host species (Garbarczyk 1982).
A city can also be considered as a dry island. Its stony environment causes that the air of a city is dryer than in its surroundings (Blume 1993). Generally the soil is also dryer in a city, despite the fact that it rains more often (Kuttler 1993). It appears that the proportion of hygrophilic species of several groups of insects will be lower inside a city than in its surroundings (Ba?kowska 1981a, Pisarski and Kulesza 1982, Chudzicka and Skibi?ska 1994). A shift from hygrophilic species to dry-tolerant species from the outskirts of a city to the centre is ascertained in Warsaw for several groups of animals: worms (Kasprzak 1986), spiders (Klausnitzer 1987), harvest-spiders (Czechowski et al. 1981b), springtails (Sterzy?ska 1982, 1987), carabid beetles (Czechowski 1981a, 1982), lady beetles (Czechowska and Bielawski 1981), flies (Górska 1981), scorpion flies (Czechowska 1982), ants (Pisarski and Czechowski 1978, Pisarski 1982b) and parasitic wasps (Garbarczyk 1982). However, it appears that dry- tolerant species with a narrow ecological amplitude occur less often in the city of Warsaw (Pisarski and Kulesza 1982). Soil and water in a city is enriched with nutrients from garbage and dung. Garbage from vegetable or animal origin is easy attainable in a city and attracts many animal species (Spirn 1984). It explains the great proportion of animals here which live on organic garbage (Trojan et al. 1982, Ba?kowska et al. 1985). The combination of a relative warm microclimate and the availability of an abundance of food makes a city attractive for many insect species (Frankie and Ehler 1978). A lot of them are most common in urban environments and are therefore known as synanthropic species. The number of species which live from organic substances (detritus) decreases to the centre of the city, e.g. mites (Niedba?a et al. 1982). Part of these detritus-eating species, which can live up in green areas in the centre occur here in great densities. These species are synanthropic and have a wide distributional range (J?dryczkowski 1981). Several carrion-beetles and carrion-eating flesh-flies (Calliphoridae) belong to this category (Klausnitzer 1989, Trojan et al. 1982, respectively). Species which larvae live from dung, like dung-flies, profit from the presence of dogs. Especially the dung-fly Scataphaga stercoraria can be very common locally (Draber-Mo?ko 1981a). The proportion of synanthropic species of many animal groups increases from the outskirts of Warsaw to the centre, as found in dung-flies, midges, sow-bugs, millipedes and spiders (Górska 1981,Wegner 1982, J?dryczkowski 1981, J?dryczkowski 1982, Krzy?anowska et al. 1981, respectively). The high density of some mammal and
bird species in the city is also explained by the availability of plenty
of food (Luniak 1980, 1981, 1983; Sukopp 1990). Especially garbage-, seed-
and fruit-eaters can find more than sufficient food. A city gives shelter
to relatively more omnivorous birds, like the magpie, jackdaw, rook and
starling. Some species, like the pigeon, have adapted their menu to city
life and changed from seed-eaters to omnivorous birds (Klausnitzer 1989).
A few birds of prey, like the peregrone falcon and kestrel, take
advantage of this situation and catch more birds in the city than in its
surroundings (Klausnitzer 1989).
In general air pollution increases towards the centre of a city (e.g. for Warsaw: see Wyrwick? 1995). This becomes evident by considering the distribution of Lichen species which are sensitive to SO2 (van Dam et al. 1986, Sukopp and Werner 1983). Air pollution has also a negative effect on the vitality of trees. Decreasing vitality of trees may lead to an increase of plant sucking insects, like cicadas (Chudzicka et al.1979, Chudzicka 1986ab) and aphids (Pisarski and Czechowski 1978). In contrast, leaf-eating insects, like caterpillars, beetles and sawflies, occur generally in low densities in the centre of a city (Pisarski 1976, Chudzicka and Skibi?ska 1994). The effects of acid deposition on the soil fauna will be strongest in sandy areas with a low content of organic material. Increasing acidification of the soil may result in a strong decrease of earthworms and certain soil-arthropods, like springtails and click-beetles (Sterzy?ska 1987, Nowakowski 1986). The quality of the soil can also be influenced by trampling. Intensive trampling causes soil condensation, which affect adversely the number of micro-arthropod species (Garay and Nataf 1982). In general disturbance of green areas increases also nearer to the centre of a city. Consequently, species which are sensitive to this factor will decrease in numbers and may in the end disappear from disturbed areas e.g. soil- and bush breeding birds may disappear from green areas which are visited intensively and birds which can sing only softly will disappear from green areas near noisy roads (Luniak 1981, 1983). However, it seems that traffic noise is not a nuisance for most bird species: the composition of the breeding population did not change significantly over a distance of 1-400 m from a busy road in Warsaw (Luniak 1981). A city is also a light-island. Illumination can disturb animals which are active at night: it can hamper their foraging (e.g. owls) and their orientation (e.g. night butterflies and migrating amphibians). Disorientation may increase the risk to become a road victim. Consequently, local populations of sensitive species may eventually disappear (de Moolenaar et al. 1997). The urban environmental factors,
mentioned above, can explain the decrease in number of species of animals
and plants from the outskirts of a city to the centre, as is apparent from
inventories which are carried out in several cities, like in Warsaw (Pisarski
1982a, among others). However, the decrease of number of species
to the centre can also be explained by the smaller size and more isolated
position of habitat patches of many species, due to the increasing density
of buildings and roads. The question arises in what respect urban biodiversity
will be affected by these factors.
Urban biodiversity Urban biodiversity can be kept on
a high level by maintaining characteristic ecotope types (ecosystems) and
species. The maintenance of biodiversity should not be aimed at maximizing
the number of species, because many exotic species are introduced by man,
intentionally or accidentally. In general green urban areas, like parks
and gardens, differ from their potential natural vegetation. Amply half
of higher plant species which occur in the centre of a city are introduced
by man (Jackowiak 1994). Part of these species which established recently
can only live up to a couple of years in this new environment. No efforts
are required to keep them. The same can be said of weedy invaders.
The number of characteristic species which occur in a city will be greater the more ecosystems are present. The same applies to a more restricted area, like a park. The larger the park the easier it is to maintain different ecotypes: e.g. woodland, shrub, grassland, marshland and water. Not only stenotopic species, which are bound to a specific ecotype, can profit from ecotope variability, but also species which need a combination of biotopes for their living and reproduction, e.g. amphibians hibernate in woodland and reproduce in water. Variation of composition and structure of the vegetation within a specific ecotype contributes also to species diversity, e.g. some species, which can be found in deciduous forests cannot be found in coniferous forests, and vice versa and some species which can be found in dense woodland cannot be found in open woodland, and vice versa. In general only part of a biotope type is suitable as habitat for a species. For example the nuthatch needs old deciduous trees (with holes) for building its nest, while the willow-warbler, as a ground breeder, prefers young open parts of a deciduous forest with some undergrowth. For maintaining biodiversity at a
high level it will be necessary that areas with a specific biotope type
(ecosystem) should not become isolated to the extent that many indigenous
species can no longer exchange individuals. Therefore, several ecosystems
should be maintained in a proper setting of a network which makes the exchange
of individuals of characteristic species possible.
Green infrastructure Water ways, like the river Vistula, as well as roads and railways to other towns, are important determining factors for the way a city expands. As a result the built- up area of Warsaw has a lobate outline: the areas in between the built-up offshoots are relatively green. These so-called green wedges can function as a corridor for the dispersal of species from the outskirts to the centre of the city. However, there are many barriers to pass, like roads and railways. This is especially difficult (or impossible) for species which cannot fly. On the other hand many of these species can disperse lengthwise along verges of roads and railways. In this respect broad verges, if properly managed, can function as habitat and corridor for characteristic species. Such verges are part of the green infrastructure of the city, together with nature reserves, parks, allotment gardens and cemeteries. A proper urban green infrastructure contributes to the biodiversity of the city and makes it more attractive as a residence. Its contribution to the species richness depends upon the size, quality and configuration of green areas. The presence of some relief contribute to biotope differences, e.g. the west side of the old river bed is marked by an escarpment, along which some parks are situated on the transition of the low river terrace (with wet areas) to the 10 – 25 m high terrace of the old river bed: an ideal situation for the establishment of a green corridor (Wolski 1999, Szulczewska and Kaliszuk 2003). Part of the green infrastructure of Warsaw is connected with a blue infrastructure. The Vistula river with its adjacent riparian forest, the streams which flow into the river and the oxbow lakes, which are remnants of the old river bed, are all parts of its blue infrastructure. Species which are living in (or near) water have a higher survival probability if these waters are part of a habitat network, that is to say if they are not only suitable as habitat, but if they can also function as dispersal corridor or stepping stone. For estimating the survival probability
of species a distribution map of their habitat is needed. For good dispersing
species the total habitat area will be most important for their survival,
while for poor dispersing species number, size and configuration of habitat
patches will be crucial. In the next paragraphs area, connectivity and
quality of habitat patches will be discussed, successively.
On theoretical grounds it is assumed that the minimum viable bird population will be at least 20 breeding pairs (Verboom et al. 1997). This implies that a viable population of tawny owls should have the disposal of a foraging area of 400 – 800 ha. This cannot be realized in one large habitat area, but only in a network of habitat patches (woodlots), between which exchange of individuals is possible. Local populations should become extinct one after the other if there should be no immigration of individuals from other patches. In the case of the tawny owl immigration is still possible if the distance between habitat patches is less than 15 km (Mooij 1982). The regional persistence of the species depends upon the proportion between the extinction of local populations and the (re)colonization of empty habitat patches. The turnover of local populations depends upon the size and connectivity of the habitat patches, the dispersal capacity of the species and the permeability of the city landscape for the species. It is argued that in a situation where the habitat of a species is fragmented the total habitat area needed for the persistence of such a metapopulation should be much greater than in a situation where its habitat is not fragmented (Verboom et al. 1997). This will be especially the case for species with a poor dispersal capacity. Stenotopic species, which are fastidious about habitat quality, are also sensitive to habitat reduction, because a decreasing proportion between area and perimeter may affect its habitat quality more severely, due to increasing deteriorating effects from its surroundings. On the basis of distribution data of stenotopic forest carabids it is concluded that a patch of woodland with a diameter of less than 80 m is unsuitable for them (Mader and Mühlenberg 1980). This may mainly be due to negative influences from the surroundings on the microclimate of the patch, although other factors may play a role as well, like pollution, disturbance and dunging (by dogs). In Warsaw the butterfly Maculinea teleius is threatened by habitat reduction and even by disappearance of its habitat. The caterpillars of this rare butterfly are dependent on Sanguisorba officinalis as food-plant and on nests of Myrmica-ants for their hibernation. The butterfly is protected by law, but will disappear from Warsaw as the plan to build houses on that location will be realized. A reduction of habitat area will generally result in a decrease of stenotopic species, while eurytopic species, with a wide ecological amplitude, will become more dominant. In Warsaw it is found that the species composition of different invertebrate groups will shift to more common species, the smaller the area of the habitat, as found in carabid beetles (Czechowski 1981a), cockchafers (Kubicka 1981), leafbeetles (W?sowska 1981, 1986), noctuids (Wegner 1982, Winiarska 1986), bees (Barnaszak 1982), wasps (Skibi?ska 1978, 1982a, 1986b), digger wasps (Skibi?ska 1982b, 1986ac), parasite wasps (Sawoniewicz 1982, 1986), ants (Pisarski 1982b), flies (Ba?kowska 1981abcd, Draber-Mo?sko 1981acd, Durska 1981, Trojan 1981b), midges (Wegner 1982) and harvest-spiders (Czechowski 1981b). The conclusion is that habitat loss, as a result of the reduction or disappearance of green areas, will lead to the extinction of local populations of species and may even lead to their disappearance from the city. Habitat connectivity Local populations of species fluctuate
in numbers. They will eventually become extinct, unless they can be rescued
by immigrating individuals from other habitat patches (Brown and Kodric-Brown
1977, among others). The immigration rate of a habitat patch depends upon
the reproduction and dispersal capacity of the species, the distance to
the nearest occupied habitat patches and the resistance of the area in
between (habitat connectivity). A species can persist in a city as long
as the extinction of its local populations can be compensated by the colonisation
of unoccupied habitat patches. This may be possible if the patches are
connected. In other words if they are part of a habitat network.
For animals which can only walk or
creep, roads, parking places and water ways are important dispersal barriers.
Nevertheless, small mammals, like the house-mouse, the longtailed fieldmouse
and the brown rat can reach all parts of a city (Klausnitzer 1989, Sukopp
1990, Szacki et al. 1994). Even larger mammals, like the fox and the beech
marten, penetrate a city sometimes very deeply at night (Harris 1986, Klausnitzer
1989, Sukopp 1990). These species disperse best in city quarters with large,
joining gardens and in green strips with bushes. Also mice and voles, which
avoid paved roads (Mader and Pauritsch 1981), use such green structures
during dispersal (Liro and Szacki 1987, 1994; Szacki et al. 1994). A subterranean
species , like the mole, can disperse via road verges and joining gardens,
although structure and food supply of the soil is decisive if he will use
them (Haeck 1969). Moles avoid built-up and paved areas. Young ones may
cross an asphalt road now and then, but often they cannot reach isolated
gardens and parks (Sukopp 1990). Hedgehogs can disperse in a city via road
verges, parks and gardens. They cross traffic roads at night, which causes
many traffic victims. Consequently, they cannot reach isolated areas (Klausnitzer
1989). That holds also for the squirrel, which is active in daytime and
will cross a traffic road rarely (Klausnitzer 1989, Sukopp 1990). The result
is that less mammal species occur in the centre than in the outskirts of
a city.
Not only nature reserves and parks can be used by species as dispersal corridors and/or as habitat, where they can reproduce, but also allotment gardens, cemeteries, road and railway verges. For most invertebrate species which disperse over short distances, the availability of habitat will be conditional for their occurrence in such anthropogenic ecotopes. Therefore it is important to pay also attention to the ecological quality of these green areas. Habitat quality Many ecotypes (biotopes) can be distinguished in a city, ranging from artificial to natural. The quality of natural biotopes generally decreases to the centre of a city, but this does not hold for artificial biotopes: some species occur mainly in the city centre, where old buildings are present, e.g. swift and kestrel (Luniak et al. 2001). Biotope quality can be maintained (or improved) by proper planning and management. A few biotope types will be discussed: woodland, grassland, fallow land and water. Woodlands are relatively stable:
they have a rather constant microclimate. In such an environment stenotopic
forest species can be expected. However, stability of woodlots may be less
in the centre of a city than in the outskirts, where it will be less intensively
used and managed. Less forest species were found in parks of Warsaw than
in its surroundings in case of carabid beetles (Czechowski 1979, 1982),
weevils (Cholewicka 1981), rove beetles (Klausnitzer 1989), julides (J?dryczkowski
1982), spiders (Krzy?anowska et al. 1981), springtails (Sterzy?ska 1982)
and several other groups. This is not only due to the more isolated position
of woodlots for species with a poor dispersal capacity, but also to disturbance
(e.g. by dogs) and intensive management of urban woodland. For example,
raking leaf litter intensively has a serious reducing effect on the species
richness of snails (Klausnitzer 1987, 1989), worms (Kasprzak 1986), julides
(J?dryczkowski 1982), spiders (Krzy?anowska et al. 1981), cicada (Chudzicka
1982, 1986), carabid beetles (Czechowski 1982) and parasite wasps (Garbarczyk
1982). Locally the species spectrum of carabid beetles, click beetles and
springtails shifted towards underground living species, due to intensive
measures of management (Czechowski 1982, Nowakowski 1982, Sterzy?ska 1982,
1987), while the spectrum of ant species shifted towards species which
live in trees (Czechowski et al. 1990). The poverty of invertebrates on
such localities is of great disadvantageous for many mammal species and
birds (Dickman 1987, Mulsow 1982, respectively; see Fig. 1). For example
the robin breeds only in parks of Warsaw where leaf litter was not raked
and birds which breed on the ground or in bushes prefer localities where
management is lacking or very extensive (Luniak 1981).
Grasslands are often intensively
managed, especially in parks for recreation activities. Consequently, these
grasslands are poor in species. This holds also for grasslands between
buildings. Often these grasslands are mowed frequently, while raking leaf
litter will impoverish the fauna even more.
Planning and management Biodiversity in a city can be kept at a high level by implementing a sustainable green infrastructure. In Warsaw the concept of an Urban Natural System was presented in the Warsaw Master Plan of 1992. The aim was to protect the most valuable green areas. However, a comparison with the adjusted Plan of 2001 it reveals that some green areas vanished (Szulczewska and Kaliszuk 2003). Protection of green areas on the level of the city should be effective, but as remarked in Warsaw Master Plans (1990, p.32): The problem of uncontrolled development will be particularly difficult to solve correctly in Warsaw, where the districts have an important voice in the matter. Biotope patches which will become smaller and more isolated will loose species, which are sensitive for habitat fragmentation. In practice it will be possible to increase habitat area of some target species by adjusting management. For example if the marshland of Zakole Wawerskie would be managed properly, then the habitat area of many characteristic and rare species will increase, and hence their survival probability. In many cases it will also be possible to increase the survival probability of species by improving the connectivity with other habitat patches by planning and implementing corridors. The river Vistula and its banks, which are partly covered by riparian forest and flower-rich meadows, is a good example of such a blue-green corridor. If properly managed it will have an important habitat and dispersal function for water and marshland species. Another example is the planned system of linear parks along the escarpment of the old riverbed in southern Warsaw (Scarpa Ursynówska). It is also desirable to plan green corridors along streams, although such ideas are not always implemented during the execution of a development plan (for Bia?o??ka Dworska, see: Ba?kowska et al. 1985). In practice the concept of an Urban Natural System will be part of “a coherent package of objectives, principles and priorities for the desired quality of green areas in the public domain“ (Meeus 1989). The question is how the implementation of an ecological network of habitats, corridors and stepping stones for the survival of species can be integrated in the planning of urban projects. In many cases it will be possible to link the ecological network with a recreational network, a traffic network and a network for proper water management (Tjallingii 2003). For example broad verges along roads and railways can be used as corridors by cyclists and/or pedestrians, as well as by animal species (Mabelis 2004, Figs. 2, 3). Another example: wasteland can be used by plants and animals as habitat and/or stepping stone, but also by children to play and experience nature. The value of wasteland for these functions is generally underestimated. In order to keep biodiversity at
a high level monitoring of nature quality is necessary. A monitoring programme
can be part of an Ecological Policy Plan of the city. Monitoring has two
functions: to determine if policy targets concerning biodiversity are attained
(evaluating or controlling function) and to signalize unexpected changes
in environmental quality (signalizing function). For both functions it
will be necessary to relate measured data to possible causes of changes
in environmental quality (diagnostic capacity). So it is not enough to
collect data, but they should be analysed and interpreted (Fig 4). For
the interpretation it is desirable to measure also possible causal factors,
like pollution and recreation pressure. The following steps should be described
in a monitoring programme:
For the registration of trends in
environmental quality a map on which the distribution of different ecosystems
(biotope types) is given will be helpful for measuring the total area per
biotope type, as well as the size and configuration of biotope patches.
Changes in the size and configuration
of green areas will affect the liveability of a city for citizens and its
viability for plants and animals. Many green areas in Warsaw have no destination
at the moment. For city planners it will be useful to make predictions
about the consequences of the expansion of built-up areas for the
survival of species, because it will make it easier to make a choice between
alternatives. For this aim species can be selected which are easily detectable,
which differ in survival strategies and which belong to different functional
groups (e.g. reducers, herbivores, carnivores).
References Bakowska, R. 1981a. Dolichopodidae
(Diptera) of Warsaw and Mazovia. Memora-bilia Zoologica 35: 33-45.
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Illustrations:
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Fig. 1: Raking leaf litter intensively impoverishes
flora and fauna (Warsaw)Fig. 2: Flower-rich road verge in Warsaw as habitat
and foraging area for insectsFig. 3: Railway verge as habitat and corridor for species
(Warsaw - Rakowiec)Fig. 4: Controlling function of a monitoring system Back to top/ terug naar begin