Introduction
Old trees is the habitat for a diverse fauna and flora. A large and well-known proportion of this fauna are beetles (Coleoptera) (Warren and Key
1991), among which are many red-listed or threatened species (Ranius and Jansson
2000; Speight
1989). Parkland, which often contains old trees, may therefore be a valuable resource for the conservation of these species (Carpaneto et al.
2010; Ehnström and Waldén
1986). Parkland, however, differs from other sites with old trees, as it is intensively managed in order to achieve the aesthetic effect of a large, tidy garden. Such intensive management is likely to be detrimental to saproxylic insects as it may often involve the removal of dead wood from the ground and tree crowns. Furthermore, old parks usually contain few bushes and small trees that might contribute to the habitat pool of dead wood. Nevertheless, studies conducted in parks and avenues have shown that they are used by threatened species (Gerell
2000; Jonsell
2004,
2008; Oleksa et al.
2006; Sörensson
2008). However, no quantitative comparisons between parks and other sites exist; this paper therefore aims to measure how parkland and more natural sites compare in their conservation value for saproxylic beetles.
The fauna of ancient trees is threatened because these trees have become increasingly rare in large parts of Europe, especially in the west (Emanuelsson
2009). Until relatively recently, old trees were widely distributed over large parts of the European landscape and were traditionally coppiced or pollarded at regular intervals to harvest wood for construction, fuel and for manufacturing wooden tools, and to provide fodder for cattle. Most of these woodlands have now been replaced by ‘high forests’ for timber production, or with modern agricultural lands. In Sweden, this decline of old trees is well documented for oak (Eliasson and Nilsson
2002). The ancient trees which remain until today were most often growing on land owned by the nobility, who could afford to keep them in parks or other semi-natural land. A century ago this land consisted of wooded meadows used for grazing, hay production and/or hunting. Today some of these areas are still kept open by grazing or they have regrown with young trees while the rest have been transformed to land without old trees. Land where the old trees still remain are highly prioritised in conservation work with protection and restoration.
In Europe, parks were often established around manor houses in the late 1600s or in the 1700s. Avenues of trees were an important feature of parks, with lime (
Tilia spp.) being the most popular species at that time (Bengtsson
2005; Sernander
1926). In most of these old parks, at least in Sweden, some 300-year old trees still remain from the original plantings (Bengtsson
2005). A number of the original trees have died, but these have usually been individually replaced, so creating a continuous supply of trees that might grow into old age. As manor houses are relatively abundant in the countryside of the region where the present study was conducted, their parks probably harbour a considerable proportion of all the ancient trees present on a landscape-scale.
The tree species studied in this paper is lime (
Tilia spp.), which hosts fewer saproxylic beetle species than, e.g. oak (Palm
1959). Compared to most other deciduous tree species, however, lime has a comparatively large assemblage of specialised saproxylic beetle species (Ehnström
2006; Palm
1959; Warren and Key
1991). But in general host specific differences in the fauna of ancient trees are not large because associated species are not usually confined to a single host species (Warren and Key
1991). Instead, the unique structures, such as hollows, dead parts of the trunk, dead branches, etc. are the important features. Because old lime trees are so frequent in parks, they might constitute an important proportion of habitat available at a landscape-scale, and so contribute to the long-term persistence of populations of saproxylic beetles.
The questions addressed in the present paper are:
(1)
Can park trees host a saproxylic beetle fauna as diverse as that found in trees of more natural stands?
(2)
Is there a difference if the natural sites are open grazed or re-grown?
Discussion
For saproxylic beetle species living in tree hollows and for red-listed saproxylic beetles species, species numbers did not differ between parks and the more natural sites. Also for species associated with wood and bark rather high numbers were found in the ‘Park’ sites, but their numbers were significantly lower than in the ‘Open’ sites. This shows that the old trees in parks harbour a rich fauna in spite of the more intensive management. The removal of wood from parks probably explains the significantly lower number of species associated with wood and bark. However, even among them, the red-listed species showed no such pattern, indicating that they could be living within the dead wood still attached to the living parts of old park trees.
Although the ordination revealed the species composition in ‘Park’ sites to be significantly different from other sites, few species discriminated between the two types of sites. The nine species showing association to ‘Park’ versus ‘non-Park’ is just one more than the number expected from the mass-significance effect (5% of the 164 tested species = 8.2). Expanding the analysis to compare all three ‘types’ gave 16 associated species, which is still marginally more than expected from mass-significance. Thus, the analysis shows that parks can be useful sites for almost all the species encountered in this study.
Sverdrup-Thygeson et al. (
2010) found parks to be species-rich sites for the saproxylic beetle fauna of hollow oaks. However, their definitions differed from those adopted in the present study since their ‘Park’ would have included the sites defined as ‘Open’ in this paper. Using a similar definition to that used in the present study, they found ‘Open’ sites to have the same numbers of red-listed species as ‘forest’ sites. However, their ‘Open’ sites had a higher proportion of species associated with hollows, which agrees with results from a study of Swedish oaks (Ranius and Jansson
2000). This suggests that regarding the hollow–dwelling species in the present study, the insignificantly higher numbers found in ‘Open’ sites compared to the ‘Re-grown’ sites was more likely to be due to the low power of the analysis, rather than any lack of a real difference. Conversely, since lime is a shade-tolerant tree it might be expected to harbour a fauna comprising fewer species adapted to sun-exposed habitats (Gärdenfors and Baranowski
1992). However, most species associated with hollows are not specific to certain tree species, and there are probably more species on lime that prefer exposed habitats than prefer shaded habitats. In the parks, the positive effect of openness seems to compensate for the negative effects from the removal of dead wood.
A problem with comparing sun-exposed sites to more shaded is that the catchability of beetles in open traps might be higher in sun-exposed sites as insect activity often is larger at higher temperature. Usually this effect is not considered at all (e.g. Sverdrup-Thygeson et al.
2010) or just assumed to be low with no reference to data (e.g. Ranius and Jansson
2000). However, Wikars et al. (
2005) found that window trapping and methods sampling directly from the wood gave similar relations in species numbers in sun-exposed and shaded environments. Thus, the assumption of low difference in catchability seems true, but more studies would be valuable and could easily be conducted by analysing already collected data.
In this paper no sites were included that could be categorised as forest because old lime trees in the region almost always grow on sites that were part of an agricultural landscape a 100 years ago, i.e. wooded meadows. For trees that exhibit traces of having been pollarded, any other situation is extremely unlikely. But trees with no such traces might originally have grown in sites that resembled forest, but which were grazed by cattle, so keeping them more open than forests are today (Emanuelsson
2009).
Generally, tree circumference explained little in this study. This is somewhat surprising as tree diameter has previously been shown to be positively correlated with the number of species (Grove
2002; Ranius and Jansson
2000; Sverdrup-Thygeson et al.
2010). However, in the present study, the trap catches and the circumferences are estimates relevant on stand scale rather than on the scale of individual trees. Therefore, other variables might have confounded the results. Furthermore, all sites were characterised by trees that had reached a size and age defining them as ancient, and the degree of ancientness may be more important than diameter itself. Pollarding slows down growth and because of that, thin trunks may be ancient trees. In oaks, 50% of trees form hollows by about 250 years of age (Ranius et al.
2009). For lime trees, this age is probably lower, as lime rots faster than oak and especially so in pollarded trees as the formation of hollows is enhanced where branches are shed. However, hollowness need not imply a rich fauna if the trees are too young, as seen in the case of 80-year old hollow limes in the park at Drottningholm, which had fewer species, especially red-listed species, than the old limes in the same park (Jonsell
2008).
The amount of habitat, measured as number of hollow lime trees on each site (No. of trees), had significant relationship to species number for all wood and bark living species, and it was negative. This lack of relation, or relation opposite to what should be expected, could be due to that the variables no. of trees and type were confounded with somewhat more trees in parks than in the other type of sites (2.6 compared to 1.9 for the two others). Also problems with quantifying this variable may contribute. First the data collected for each locality had several uncertainties in itself (see “
Materials and methods”). The numbers obtained also give just the present situation, totally disregarding the history of the site. In addition to that, the definition of where the borders for a locality should be drawn is also problematic. Most of these sites are found in regions where old hollow trees may occur here and there. Data on suitable trees for the whole landscape with estimates of connectivity related to distance to each of these occurrences should probably be more explanatory (Ranius et al.
2010). Such an analysis would probably suggest that the rich saproxylic beetle fauna on several sites in the Mälaren area is due to a dense patchwork of sites. The number of sites is high, there is a high connectivity between them, several sites are large and the individual trees in them are often a high quality habitat, all factors that contribute to a sustainable metapopulation system (Hanski
1994; Ranius
2007). Two practical conclusions from this is that every park with old trees constitutes a site that contributes to the sustainability of the metapopulation and that the number and quality of trees within each park also contributes to this.
The north–south coordinate was a strong explanatory variable both for species numbers and species composition. This is not surprising because, compared to the sites north of the lake, the area around lake Mälaren is both climatically favourable (Raab and Vedin
1995) and has a high density of sites with old trees. Mälaren has been identified as a diversity hot-spot for saproxylic beetles (Ehnström and Waldén
1986), with the western part of Mälaren regarded as being especially species-rich. This was only weakly supported by the results of the present study, as the variable RT90E (west–east coordinate) had low explanatory power.
Practical implications
The high conservation value of parks for saproxylic insects shown in this study is dependent on the retention of old trees. Thus, the total rejuvenation of trees, which is considered in some parks, would be fatal to the resident fauna. However, all trees will sooner or later die, or they have to be removed for safety or aesthetic reasons. If they individually and continuously are replaced when they die there will be a continuous supply of new trees growing into the ancient-tree age class which in turn means a continuous supply of suitable habitat for the saproxylic insects.
On a short term a good measure is to retain trees, or parts of trees, that are cut or fallen in a “tree-graveyard” situated in a remote part of the park, where it does not conflict with the aesthetic values. Such graveyards is both a chance for insects to finalise their development and a habitat patch that can be colonised (Aulén and Franc
2008). However, compared to the management aiming at a long term continuous supply of old trees, this is of minor importance, both because its’ short term effect and because most of the valuable contributions to the graveyard emanate from the old trees.
As almost all lime trees in parks, and many lime trees in the more natural sites, were originally pollarded, they are at risk of breaking apart when the shoots from the last pollarding are allowed to grow into large trees. This was observed on several of the sites in this study. The risk of breakage is especially great in re-grown sites where the closer canopy gives less light to the trees, which in turn decreases the production of carbohydrates needed for building a stable trunk. For keeping these old trees alive it is important to continue pollarding. However, old trees that has not been managed for a long time need careful treatment when management is resumed (Slotte
1997; Wisenfield
1995). A strong reduction of the crown by cutting all large branches may be fatal. As pollarding is an expensive measure, it is important that it should only be done on sites where there is the potential to retain the associated fauna and flora, i.e. where one can forecast a continuous supply of old trees in the future. Most of the parks in the present study do have this potential due to the continuous replacement of trees that die.
This study has shown that old trees in parks are a very valuable substrate for saproxylic beetles and thus, they may contribute to the long-term survival of these species. In the management of the parks, however, conservation of the biological values can be in conflict with measures undertaken to enhance or preserve historical and cultural values (Jonsell
2008) or measures for public safety (=risk for falling parts of trees) (Carpaneto et al.
2010). Such conflicts often arise because park managers are not always aware of the high biological value of elements within their domain and vice versa. The biologists may not reflect over the heavy cultural influence on the habitat of “their” species. Thus, effective communication between those with different interests is important in order to share knowledge about different sets of values so that both the cultural and biological value of a site can be maintained for the benefit of all.