The network between birds and preferred tree species
As we hypothesized, the bird-tree preference connections in our study revealed an unfragmented network with key members of both bird and tree species.
As we recorded, great spotted woodpeckers used a wide range of tree species, being the most generalist woodpecker species in the Western Palearctic (Gorman
2004; Ónodi et al.
2021; Stański et al.
2020,
2021a). In a study on its foraging behavior in an oak-lime-hornbeam forests of Białowieża National Park, they also preferred a wide range of tree species,
e.g., aspen (
Populus tremula), small-leaved lime (
Tilia cordata), Norway maple (
Acer platanoides), common hornbeam and pedunculate oaks (Stański et al.
2020).
Similarly to other studies (
e.g., Pasinelli
2000 Stański et al.
2021b), middle spotted woodpeckers highly preferred pedunculate oaks, as old oak trees with decaying parts can have a variety of foraging sources for this woodpecker species. In spite of this, they showed the greatest preference for girdled individuals of the invasive tree of heaven. These trees, along with non-native box elder trees (
Acer negundo), were killed by horizontal chainsaw cuts around the stems in a national park-wide effort from 2010 to 2014 (Quadt et al.
2016). Due to these efforts, a substantial proportion of snags and logs found nowadays in the National Park originate from this invasive tree species. We found no information in the literature for middle spotted woodpeckers foraging on
Ailanthus altissima. However, there are a few records of wood-boring insect species found in the wood of this species in Hungary that can be prey of woodpeckers (Kovács
1995,
1997; Kovács and Gebei
2021). In the oak-lime-hornbeam (
Tilio-Carpinetum) forests of the Białowieża National Park, Stański et al. (
2021b) found that apart from pedunculate oaks, middle spotted woodpeckers also preferred Norway maple (
Acer platanoides) trees, which were only found scarcely in our study area and we did not record any foraging data from this tree species. Pasinelli (
2000) also found preferential use of oaks in Swiss lowland forests characterized by oak, hornbeam and lime.
As already documented, the Eurasian nuthatch occupies a broad niche in terms of tree species use (Adamík and Kornan
2004, Korňan and Adamík
2017). This was also confirmed by our study. Treecreepers are known to use a wide range of tree species (Ceia and Ramos
2016a; Korňan and Adamík
2017), similarly to our observations. Adamík and Korňan (
2004) and Korňan and Adamík (
2017) also reported a preference for sycamore maple by both common treecreeper and Eurasian nuthatch, for which we did not find evidence.
The unique tree preference exhibited by marsh tits may perhaps be explained by its subordinate role among Parid species. Haftorn (
1993) found that marsh tits are subordinate to both great tits and also Eurasian blue tits. This may explain why we did not observe this species foraging on the scarcely found pedunculate oaks. At the same time, other studies highlight its significant role in habitat selection and winter foraging ecology of marsh tits (Broughton et al.
2014). The competitive exclusion by great tits and Eurasian blue tits relative to pedunculate oaks was also suggested by Carpenter (
2008). In a coniferous-deciduous mixed mountain forests of Slovakia, in the absence of great tits and Eurasian blue tits, but in the presence of coal tits (
Periparus ater) and crested tit (
Lophophanes cristatus), marsh tits preferred to forage on sycamore maples (Adamík et al.
2003). In our study, we could not find any preference for sycamore maples by marsh tits, but great and Eurasian blue tits did show a preference for this particular tree species. This preference may be due to social dominance or difference in prey availability of the same tree species at different geographical locations in different seasons (Osborne and Green
1992; Travis
1977; Unno
2002). Great tits frequently used oaks (
Quercus spp.) and sycamore maple trees as well in a study conducted in spring in parks with common ash, oak, sycamore maple, beech, elder, hawthorn, hazel, blackthorn (
Prunus spinoza) and willow (
Salix spp.) trees in Oxford (Franzreb
1984). In that study, Eurasian blue tits foraged most frequently on common ash and oak trees, while in our study they did not use oaks but preferred common ash trees. Marsh tits in Franzreb’s (
1984) study foraged mostly on black elderberry and common ash trees; we also found a preference for the latter tree species. Rolando (
1982) studied the autumn–winter foraging behavior of these three particular tit species in parks in the Po River valley, with dominant tree species such as pedunculate oak, common ash, and poplars. He found that marsh tits foraged on oak trees the least among tit species and foraged the most on hazel. We also found a preference for hazel.
We found the European white elm to be the most connected and essential key species for this bird-tree network, as all bird species showed preferences for this particular tree species. The European white elm is symbolic of temperate European oak-ash-elm floodplain forests. As a late successional tree species, it is susceptible to the drier climate caused by climate change, but also to the effects of river regulations with deeper riverbeds and thus a lower groundwater table (Allen et al.
2010; Hemery et al.
2010; Venturas et al.
2014). Studies on bark-foraging birds using elm trees are rare, especially in the case of the European white elm. In North America, downy woodpeckers (
Dryobates pubescens) prefer dead American elm trees (
Ulmus americana) in winter (Jackson
1970). In a study on the tree preferences of tree-foraging insectivorous bird species in a primeval beech-fir forests of Slovakia, the wych elm (
U. glabra) was most preferred by species also considered in our study, namely Eurasian nuthatch, Eurasian treecreeper, and marsh tit (Korňan and Adamík
2017). According to Holmes and Robinson (
1981) and Holmes and Schultz (
1988), elms, besides other tree species such as willows and sycamores (
Platanus spp.), are disturbance-dependent species that grow fast and generally have lower defenses against herbivory and therefore have higher abundance and biomass of insects. As elms have relatively low population densities but have a significant role in bird foraging, it is crucial to take the conservation of the European white elm into consideration among other secondary tree species in floodplain forest habitats (Venturas et al.
2014).
The high importance of
Quercus robur is equivocal with other studies, as oak species provide a high diversity of microhabitats with essential food sources for bark-foraging birds (Domokos and Cristea
2014; Ónodi et al.
2022; Proença et al.
2010; Robles et al.
2011; Stański et al.
2021b).
Tree characteristics
In our study, species that are particularly specialized on searching for insects on and under bark showed a clear preference for decayed and dead trees. This was also the case in the study of Korňan and Adamík (
2017), which was conducted in a primeval mountain mixed forest in Slovakia, where among the numerous tree-foraging bird species, only the Eurasian nuthatch and the studied two woodpecker species, namely the three-toed woodpecker (
Picoides tridactylus) and the white-backed woodpecker (
Dendrocopos leucotos), showed preference for dead trees. Also, in our study, besides the importance of tree species identity, there was at least a weak preference for more decayed trees in both woodpecker species compared to other bark-foraging birds. Excluding treecreepers, all species studied showed a preference for larger DBH.
In this study, apart from treecreepers, all other bird species preferred trees with a larger diameter and a rougher bark structure. In an earlier study by Ónodi et al. (
2021) on the winter intersexual foraging segregation of great spotted woodpeckers in softwood willow-poplar forests in Hungary with a high presence of invasive non-native tree species, such as green ash (
Fraxinus pennsylvanica) and box elder (
Acer negundo), this woodpecker species preferred the rougher bark willow and poplar trees with larger DBH, compared to the invasive tree species with smoother bark surfaces. In those forests, willow and poplar species provided more foraging sources compared to non-native trees. In contrast, the floodplain forests of the Donau-Auen National Park have a high number of native species and, therefore, a wider variety of food sources. Great spotted woodpeckers also chose the larger-diameter trees with rough bark structures. Similarly to our results, in the highly diverse oak-lime-hornbeam stands of Białowieża National Park in Poland, Stański et al. (
2020) found a preference of this woodpecker species for more decayed foraging trees with a greater DBH. Nevertheless, on the level of bird species, we did not find a preference for decayed trees. On the preference for larger-diameter trees, Stański et al. (
2020) hypothesized that trees with larger trunk diameters can have a higher number of invertebrates than smaller-diameter trees.
Middle spotted woodpeckers preferred dead trees over living trees in the work of Stański et al. (
2021b) in the oak-lime-hornbeam forest stands of the Białowieża National Park in the non-breeding season. As in that particular study, middle spotted woodpeckers preferred the rough-barked pedunculate oak and Norway spruce the most, thus, bark roughness could also be an important factor for the species, as also documented by our study.
In a study on post-fledging middle spotted woodpeckers in the Cantabrian mountains of Northern Spain, in forests where oaks were the major species with a significant amount of poplars, willows, ashes, and pines, birds used only larger diameter decayed oak, willow, and poplar trees of rougher bark (Ciudad et al.
2009), similarly to our study.
In a study on the foraging of middle spotted woodpeckers made in the Taurus mountain range in Turkey during April/May (Bergner et al.
2016), in an area of mainly pollarded individuals of four oak species, among other tree characteristics, middle spotted woodpeckers preferred to forage on trees with a greater circumference and bark roughness, as we also found.
In contrast to Korňan and Adamík (
2017), who found a preference of foraging Eurasian nuthatches for dead trees in a beech-fir mixed forest in spring, we could not confirm this phenomenon in our study. This difference may be attributed to habitat and seasonal differences (Michielsen et al.
2024). In the former study, nuthatches also preferred tree species with rougher bark structure (such as sycamore maple and Norway spruce), similar to our study.
In addition to species-specific preferences for certain tree species, we found that great, Eurasian blue and marsh tits preferred to forage on trees with rougher bark. Similar results were reported by Franzreb (
1984) and Rolando (
1982) for the same tit species in similar forests consisting predominantly of pedunculate oak, common ash, and poplars. The three species almost exclusively used trees of intermediate or rough bark structure, such as common ash, sycamore maple, and pedunculate oak.
Studies by Fraticelli and Guerrieri (
1988) carried out in a Mediterranean oak-dominated forest and that conducted by Karpińska et al. (
2023) in a temperate primeval lime-oak-hornbeam and ash-alder forests showed that tree trunks and bark contributed less to the foraging properties of tit species. In the floodplain forests of our study area, all tit species were observed foraging on both the limbs and the trunk. The fact that bark roughness had a significant effect on both great and blue tits suggests that this variable plays an important role in some forest types.
According to MacFarlane and Luo (
2009), who measured bark roughness of individual trees belonging to different species in an old-growth natural hardwood forest in Michigan, USA, bark roughness of individual tree species correlated with the stem DBH, suggesting that tree aging increases the amount of microhabitats for bark-living arthropods. As more fissured bark structures can harbor more arthropod prey for bark-foraging birds (Jackson
1979), it is crucial to conserve old forest stands or apply forest management practices for retaining old, aging tree individuals in commercial forests, such as green tree retention (Machar et al.
2019).