The invasion of Lantana camara L. in Forty Mile Scrub National Park,north Queensland

Abstract Seventy-three per cent of dry rainforest in Forty Mile Scrub National Park and large areas in adjacent savanna woodland have more than 5000 individuals per ha of lantana (Lantana camara L.). Transect studies in dry rainforest and savanna woodland across varying intensities of lantana infestation show a negative correlation between the density of lantana and tree cover in rainforest. The density of pig rooting is very high in areas of the dry rainforest on deep soil that was not heavily infested with lantana. It is suggested that the digging activities of these animals may cause tree death and subsequent increased light penetration, which favours lantana. The species richness of the dry rainforest declines as the density of lantana increases. However, the saplings and seedlings and the soil seed bank of dry rainforest and savanna woodland tree species have comparable densities in heavy and light lantana infestations. The proliferation of lantana results in the build up of heavy fuel loads across the boundary of dry rainforest and savanna woodland. Recent fires have killed the canopy trees in a large area of dry rainforest within the Park. Active management of Forty Mile Scrub National Park is urgent and some initiatives are suggested.     

Holocene boundary dynamics of a northern Australian monsoon rainforest patch inferred from isotopic analysis of carbon, (14C and δ13C) and nitrogen (δ15N) in soil organic matter

Abstract Soil organic matter (SOM) was sampled from lateritic soil profiles across an abrupt eucalypt savanna–monsoon rainforest boundary on the north coast of Croker Island, northern Australia. Accelerator mass spectrometry dating revealed that SOM that had accumulated at the base of these 1.5 m profiles had a radiocarbon age of about 5000 years. The mean carbon and nitrogen stable isotope composition of SOM from 10 cm deep layers from the surface, middle and base of three monsoon rainforest soil profiles was significantly different from the means for these layers in three adjacent savanna soil profiles, suggesting the isotopic ‘footprint’ of the vegetation boundary has been stable since the mid Holocene. Although there were no obvious environmental discontinuities associated with the boundary, the monsoon rainforest was found to occur on significantly more clay rich soils than the surrounding savanna. Tiny fragments of monsoon rainforest and abandoned ‘nests’ (large earthen mounds) of the orange‐footed scrubfowl, an obligate monsoon rainforest species, occurred in the savanna, signalling that the rainforest was once more extensive. Despite episodic disturbances, such as tropical storm damage and fires, the stability of the boundary is probably maintained because clay rich soils enable monsoon rainforest tree species to grow rapidly and achieve canopy closure, thereby excluding grass and reducing the risk of fire. Conversely, slower tree growth rates, grass competition and fire on the savanna soils would impede the expansion of the rainforest although high rainfall periods with shorter dry seasons may enable rainforest trees to grow sufficiently quickly to colonize the savanna successfully.     

Mycorrhizas in the Kakadu region of tropical Australia

This research represents the first part of a study which aimed to characterize the role of mycorrhizal associations in undisturbed and disturbed habitats in the Alligator Rivers Region of the Northern Territory of Australia. This is a seasonally dry tropical region with a climate consisting of a long dry season and a monsoonal wet season. Intact soil cores were sampled from 22 sites in this region, representing eucalypt savanna woodland, wetland, rocky hill and rainforest habitats. Clover, sorghum and eucalypt seedlings were grown in these cores in bioassays to measure the inoculum potential of vesicular-arbuscular mycorrhizal (VAM) and ectomycorrhizal (ECM) fungi. Propagules of VAM fungi were concentrated in the surface horizon, and were not adversely affected by 6 months dry storage of soil. Bioassays detected VAM fungus propagules at all sites, but these were less numerous in three sites with sparse herbaceous vegetation (a shrub-dominated woodland site, a sandstone area and a disturbed gravel pit without topsoil), than in other woodland sites. Propagules of VAM fungi were particularly numerous in soil from a rainforest habitat, which had much denser plant cover than any of the savanna sites. Propagules of ECM fungi colonized eucalypt seedling roots in some cores from all sites, except two wetland areas and a disturbed area without eucalypt trees. Physical and chemical properties of soils varied between sites and some properties (texture, organic carbon, etc.) were correlated with the inoculum potential of VAM fungi.     

Vegetation,fire and soil feedbacks of dynamic boundaries between rainforest,savanna and grassland

At fine spatial scales, savanna‐rainforest‐grassland boundary dynamics are thought to be mediated by the interplay between fire, vegetation and soil feedbacks. These processes were investigated by quantifying tree species composition, the light environment, quantities and flammability of fuels, bark thickness, and soil conditions across stable and dynamic rainforest boundaries that adjoin grassland and eucalypt savanna in the highlands of the Bunya Mountains, southeast Queensland, Australia. The size class distribution of savanna and rainforest stems was indicative of the encroachment of rainforest species into savanna and grassland. Increasing dominance of rainforest trees corresponds to an increase in woody canopy cover, the dominance of litter fuels (woody debris and leaf), and decline in grass occurrence. There is marked difference in litter and grass fuel flammability and this result is largely an influence of strongly dissimilar fuel bulk densities. Relative bark thickness, a measure of stem fire resistance, was found to be generally greater in savanna species when compared to that of rainforest species, with notable exceptions being the conifers Araucaria bidwillii and Araucaria cunninghamii. A transect study of soil nutrients across one dynamic rainforest – grassland boundary indicated the mass of carbon and nitrogen, but not phosphorus, increased across the successional gradient. Soil carbon turnover time is shortest in stable rainforest, intermediate in dynamic rainforest and longest in grassland highlighting nutrient cycling differentiation. We conclude that the general absence of fire in the Bunya Mountains, due to a divergence from traditional Aboriginal burning practices, has allowed for the encroachment of fire‐sensitive rainforest species into the flammable biomes of this landscape. Rainforest invasion is likely to have reduced fire risk via changes to fuel composition and microclimatic conditions, and this feedback will be reinforced by altered nutrient cycling. The mechanics of the feedbacks here identified are discussed in terms of landscape change theory.     

Spacing patterns of an Acacia tree in the Kalahari over a 61-year period: How clumped becomes regular and vice versa

Nearest tree neighbour distances and the tree spatial formation on a large scale over time and space replicates were examined. The study was conducted in a natural savanna ecosystem in the Southern Kalahari, South Africa. Nearest tree neighbour and point pattern analysis methods were used to investigate changes in the spatial pattern of trees in two plots. Trees larger than 2 m canopy diameter were mapped. We used aerial photographs of the study area from 1940, 1964, 1984, 1993, and a satellite image from 2001 to follow two plots over time. Field work was carried out too for classification accuracy. We were able to identify and individually follow over 2400 individual trees from 1940 until 2001. Nearest neighbour analysis results indicate that dead trees were on average closer to their nearest neighbouring trees than living trees were to their neighbours. Most dead trees were on average 6 m from their nearest neighbours, while most living trees were about 20 m apart. Point pattern analysis results show a cyclical transition from clumped to random and sequentially to regular tree spacing. These transitions were not correlated across two plots. Generally, decreases in small-scale clumping coincided with periods of high mortality. Our findings show that regular, clumped, and random tree pattern can occur, pending on time, location, and scale within the location.     

Vessel feature changes as a tool for detecting annual rings in tropical trees

Radial variations in vessel features (vessel lumen area and frequency) of four tropical tree species grown in tropical savanna, monsoon, and rainforest climates were investigated to detect indistinct annual rings. Leaf and soil water potentials were measured periodically to show annual variations in water availability and their relationship to radial variations in vessel features. In addition, the accuracy of annual-ring detection was estimated using trees of known age. Radial variations in vessel features showed annual cyclicity in all three sites. The vessel feature that showed clear annual cyclicity was different among the species and sites. Furthermore, the variation pattern of vessel features in the rainforest site tended to be different between two radii within individual trees, implying low synchronicity of wood formation in the wet area. Compared with leaf and soil water potentials, vessel features varied independent of water availability in all species in the rainforest site and most species in the savanna and monsoon sites. The direct effect of low water availability on vessel features was considered only in one species in the savanna and monsoon sites. Nevertheless, the deviation of vessel feature cycles from tree age was similar and reasonably small (mostly within ±10 %) among all sites. These results indicated a circannual rhythm in vessel formation. Thus, vessel feature changes seemed to aid in detecting annual rings in trees throughout wide tropical areas; however, asynchronous wood formation must be considered in dendrochronology, especially in humid tropics.     

A comparison of foliar nutrient concentration in trees from monsoon rainforest and savanna in northern Australia

Abstract Analysis of foliar nutrient concentrations revealed that two mesophyllic monsoon rainforest trees had higher concentrations of most nutrients in leaves than eight savanna species. One of the tested monsoon rainforest species with sclerophyllous leaves had similar nutrient concentrations to the savanna tree species. There were positive or no significant correlations between live foliar nutrient concentration and the percentage of nutrients withdrawn prior to litterfall. The nutrient concentration of litter was similar for most nutrients among tree species of monsoon rainforest or savanna. The results of this study suggest that the relative fertility of surface soils of monsoon rainforest compared with savanna is not determined by contrasting nutrient cycling strategies whereby monsoon rainforest tree species enrich their soils with relatively nutrient rich leaf litter relative to savanna tree species.     

Sleeping Sites and Latrines of Spider Monkeys in Continuous and Fragmented Rainforests: Implications for Seed Dispersal and Forest Regeneration

Spider monkeys (Ateles geoffroyi) use sites composed of one or more trees for sleeping (sleeping sites and sleeping trees, respectively). Beneath these sites/trees they deposit copious amounts of dung in latrines. This behavior results in a clumped deposition pattern of seeds and nutrients that directly impacts the regeneration of tropical forests. Therefore, information on the density and spatial distribution of sleeping sites and latrines, and the characteristics (i.e., composition and structure) of sleeping trees are needed to improve our understanding of the ecological significance of spider monkeys in influencing forest composition. Moreover, since primate populations are increasingly forced to inhabit fragmented landscapes, it is important to assess if these characteristics differ between continuous and fragmented forests. We assessed this novel information from eight independent spider monkey communities in the Lacandona rainforest, Mexico: four continuous forest sites and four forest fragments. Both the density of sleeping sites and latrines did not differ between forest conditions. Latrines were uniformly distributed across sleeping sites, but the spatial distribution of sleeping sites within the areas was highly variable, being particularly clumped in forest fragments. In fact, the average inter-latrine distances were almost double in continuous forest than in fragments. Latrines were located beneath only a few tree species, and these trees were larger in diameter in continuous than fragmented forests. Because latrines may represent hotspots of seedling recruitment, our results have important ecological and conservation implications. The variation in the spatial distribution of sleeping sites across the forest indicates that spider monkeys likely create a complex seed deposition pattern in space and time. However, the use of a very few tree species for sleeping could contribute to the establishment of specific vegetation associations typical of the southeastern Mexican rainforest, such as Terminalia-Dialium, and Brosimum-Dialium.     

Fire acting as an increasing spatial autocorrelation force: Implications for pattern formation and ecological facilitation

Fire is an indissoluble component of ecosystems, however quantifying the effects of fire on vegetation is a challenging task as fire lies outside the typical experimental design attributes. A recent simulation study showed that under increased fire regimes positive tree–tree interactions were recorded (Bacelar et al., 2014). Data from experimental burning plots in an African savanna, the Kruger National Park, were collected across unburnt and annual burn plots. Indices of aggregation and spatial autocorrelation of the distribution of trees between different fire regimes were explored. Results show that the distribution of trees under fire were more clumped and exhibited higher spatial autocorrelation than in unburnt plots. In burnt plots spatial autocorrelation values were positive at finer scales and negative at coarser scales potentially indicating co-existence of facilitation and competition within the same ecosystem depending on the scale. The pattern derived here provides inference for (a) fire acting as an increasing aggregation & spatial autocorrelation force, (b) tree survival under fire regimes is potentially facilitated by forming patches of trees and (c) scale-dependent facilitation and competition coexisting within the same ecosystem with finer scale facilitation and coarser scale competition.     

Spatial pattern in the occurrence of the lichen Lobaria pulmonaria in managed and virgin boreal forests

Two populations of the lichen Lobaria pulmonaria. growing on aspens and goat willows in 12 and 20 km² study areas of boreal forest in Finland, were surveyed thoroughly to investigate the factors influencing the spatial distribution of the lichen. In one study area, where forestry has been intensive and old-growth forest is highly fragmented, L, pulmonaria was sparse and grew mostly on willows. In contrast, a large and continuous virgin forest area supported a higher incidence of L. pulmonaria. with the lichen being common on both aspens and willows. In both study areas, the distributions of aspen and willow were clumped over the scales of 100-1000 m. The spatial pattern of L pulmonaria was more clumped in the managed forest than in the virgin forest. The reduced incidence of the lichen on aspens in the managed area was attributed to a disruption of habitat continuity and small average tree size. There was no comparable reduction in the incidence on willows, probably because the willow had a very aggregated distribution in the managed area, which probably facililated local colonization of the lichen. Presence of the lichen was significantly related to size-corrected local density of aspen and willow trees as well as to spatial connectivity to neighboring lichen-occupied trees.     

Mechanisms of rainforest persistence and recruitment in frequently burnt wet tropical eucalypt forests

The role of fire in governing rainforest–eucalypt forest ecotone dynamics is of theoretical interest and has conservation management implications. Several eucalypt forests in the Wet Tropics of Australia have an endangered status due to extensive conversion to rainforest. Rainforest plants are known to survive occasional low intensity fires in the eucalypt forest ecotone. However, the ability of rainforest plants to survive frequent fires remains untested. The timing of rainforest expansion is also a subject of interest, and is generally considered to be delayed until fire has been absent for several years. We used 14 years of data collected across 13 plots in the Wet Tropics of north‐eastern Australia to test predictions regarding rainforest seedling recruitment and post‐fire regenerative capacity. The 13 plots received different numbers of fires, between zero and five, over the 14‐year study. The recruitment of new rainforest plants in the ecotone was most abundant in the initial year after fire. If this post‐fire pulse of recruitment is left undisturbed, it can facilitate the subsequent germination of additional rainforest species. The removal of grass cover, whether temporarily in the immediate post‐fire environment or once a developing rainforest mid strata shades out grasses, appears crucial to abundant rainforest recruitment. A variety of tropical rainforest species can persist under a frequent fire regime through resprouting. The difference in the mode of resprouting, between ground‐level coppicing rainforest plants and canopy resprouting eucalypt forest trees, is the critical mechanism that causes regular fire to maintain an open structure in eucalypt forests. The inability of rainforest species to maintain their height when fires fully scorch their crowns, temporarily resets the forest's open structure and delays the rainforest's ability to dominate through shading out grasses to transform the ecosystem into a closed forest.     

The spatial pattern and canopy-understory association of trees in a cool temperate,mixed forest in western Japan

The spatial distribution pattern of trees and the association between canopy and understory individuals were examined with reference to the distribution of tree crowns in a cool temperate, mixed forest in Ohdaigahara, western Japan. Line transect and contact sampling methods were used to examine the pattern over various spatial scales. These methods are useful to detect patterns over a large study area. The dominance ofChamaecyparis obtusa on steep slopes forming large patches suggested that the distribution of this species is a consequence of landslides. UnderstoryFagus crenata showed a clumped distribution, and the relative coverage of this species was larger in canopy gaps than under a closed canopy. Understory individuals ofAbies homolepis showed a positive association with canopy trees ofF. crenata but a negative association with conspecific canopy individuals. These patterns suggested thatF. crenata regenerates in canopy gaps and is replaced byA. homolepis. The dynamics of these two species are consistent with the process of gap dynamics. The effects of both small- and large-scale disturbance must be evaluated to understand the mechanisms of patch formation and the coexistence of forest tree species.     

The epiphyte vegetation of Annona glabra on Barro Colorado Island, Panama

Aim Information on the community composition, structure, and dynamics of epiphyte vegetation is scarce. A survey of the epiphytes occurring on all individuals of one particular host tree species in a well-studied neotropical research site allowed us a comparison of the epiphyte flora of this tree with the local epiphyte flora, the analysis of spatial distribution patterns and the use of these patterns as indications for changes in time. In the future, our results can be used as a baseline data-set for the direct observation of the long-term dynamics in epiphyte communities. Location The study was conducted on Barro Colorado Island (BCI), Panama. Methods We recorded all individuals of the vascular epiphytes growing on Annona glabra L., a flood-tolerant, multiple-stemmed tree, which is restricted to the shoreline of BCI. Data on tree biometrics, epiphyte species, and epiphyte abundances were collected for more than 1200 trees. Results In total, we encountered almost 15,000 epiphytic individuals in sixty-eight species, corresponding to more than one third of the entire epiphyte flora of Barro Colorado Island. The component species differed strongly in abundance: the four most important species accounted for >75% of all individuals. In most cases, the same four species were also the first to colonize a tree (=phorophyte). Colonization patterns indicated no replacement of early colonizers by late arrivals. Species richness and epiphyte abundances showed a positive correlation with the size and the density of the host trees. All species showed a highly clumped distribution and the physiognomy of epiphyte communities of individual trees was dominated either by one or several of the four most common species or by a set of frequently co-occurring tank bromeliads. Other species were dominant only in exceptional cases. Most species were always rare. A distance effect on community composition was mostly confined to a local scale with an increased similarity in the species assemblage of stems of a tree v. neighbouring trees. Main conclusions The epiphytes on a single small phorophyte species may encompass a surprisingly large proportion of the local epiphyte flora. The observations that most tree crowns are inhabited by a single or only very few species, and that all epiphyte species show highly clumped distributions suggest a predominance of very local dispersal within a tree crown, which is only infrequently interrupted by successful long-distance dispersal between crowns.     

Rainforest—eucalypt forest interactions and the relevance of the biological nomad concept

Transects across the margins of rainforests with eucalypt forests at two sites in New South Wales are described. At Girard State Forest along ten transects it was consistently found that pure rainforest occurs where there is no evidence of past burning; and that rainforest tree species appear to be invading and replacing eucalypt forest through a formerly burned ecotone area. A similar pattern was found at Barrington Tops. It is suggested that adaptations shown by species of tall open forests which enable them to regenerate rapidly after burning may have evolved initially in a fire free, prehuman rainforest environment in response to other types of disturbance.     

Tree formations in relation with soil and grasses in a dry savanna in Mali, West Africa

The relationship between trees, grass and soil in a dry savanna in Mali was investigated, to identify variables that are most relevant to assess vegetation units. A 65 ha plateau was inventoried using a systematic square grid sampling pattern. Thirteen soil or topography variables, and tree and grass characteristics were measured at each sampling point. Multivariate analysis was used to separately analyse soil, tree and grass data, and to characterize tree–grass and tree–soil relationships. Four units of soils, four units of tree formations, and four units of grass formations were identified. There was a correspondence between these groups, indicative of four vegetation units: thicket, bare land, shrub savanna and tree savanna. Soil depth and soil texture were the soil variables that best related to tree vegetation. A negative correlation was found between tree basal area and grass dry biomass. Finally, vegetation units, as identified from tree species composition, had contrasted diameter structures and densities.     

Trees improve grass quality for herbivores in African savannas

The tree–grass interactions of African savannas are mainly determined by varying rainfall patterns and soil fertility. Large savanna trees are known to modify soil nutrient conditions, but whether this has an impact on the quality of herbaceous vegetation is unclear. However, if this were the case, then the removal of trees might also affect the structure and quality of the grass layer. We studied the impact of large nitrogen- and non-nitrogen fixing trees on the sub-canopy (SC) grass layer in low- and high-rainfall areas of differing soil fertility in eastern and southern Africa. We compared the structure and nutrient levels of SC grasses with those outside the canopy. Grass leaf nitrogen and phosphorus contents beneath tree canopies were elevated at all study sites and were up to 25% higher than those outside the canopy in the site of lowest rainfall and soil fertility. Grass leaf fibre and organic matter (OM) contents were slightly enhanced beneath tree canopies. At the site of highest rainfall and soil fertility, grasses beneath the canopy had significantly lower ratios of stem:leaf biomass and dead:living leaf material. Grass species composition differed significantly, with the highly nutritious Panicum spp. being most abundant underneath tree crowns. In the two drier study sites, soil nitrogen and OM contents were enhanced by 30% beneath trees. N-fixation capacity of trees did not contribute to the improved quality of grass under the canopy. We conclude that trees improve grass quality, especially in dry savannas. In otherwise nutrient-poor savanna grasslands, the greater abundance of high-quality grass species with higher contents of N and P and favourable grass structure beneath trees could attract grazing ungulates. As these benefits may be lost with tree clearance, trees should be protected in low fertility savannas and their benefits for grazing wildlife recognised in conservation strategies.     

Plant Traits Demonstrate That Temperate and Tropical Giant Eucalypt Forests Are Ecologically Convergent with Rainforest Not Savanna

Ecological theory differentiates rainforest and open vegetation in many regions as functionally divergent alternative stable states with transitional (ecotonal) vegetation between the two forming transient unstable states. This transitional vegetation is of considerable significance, not only as a test case for theories of vegetation dynamics, but also because this type of vegetation is of major economic importance, and is home to a suite of species of conservation significance, including the world’s tallest flowering plants. We therefore created predictions of patterns in plant functional traits that would test the alternative stable states model of these systems. We measured functional traits of 128 trees and shrubs across tropical and temperate rainforest – open vegetation transitions in Australia, with giant eucalypt forests situated between these vegetation types. We analysed a set of functional traits: leaf carbon isotopes, leaf area, leaf mass per area, leaf slenderness, wood density, maximum height and bark thickness, using univariate and multivariate methods. For most traits, giant eucalypt forest was similar to rainforest, while rainforest, particularly tropical rainforest, was significantly different from the open vegetation. In multivariate analyses, tropical and temperate rainforest diverged functionally, and both segregated from open vegetation. Furthermore, the giant eucalypt forests overlapped in function with their respective rainforests. The two types of giant eucalypt forests also exhibited greater overall functional similarity to each other than to any of the open vegetation types. We conclude that tropical and temperate giant eucalypt forests are ecologically and functionally convergent. The lack of clear functional differentiation from rainforest suggests that giant eucalypt forests are unstable states within the basin of attraction of rainforest. Our results have important implications for giant eucalypt forest management.     

Fire weather risk differs across rain forest—savanna boundaries in the humid tropics of north‐eastern Australia

Alternative stable state theory has been applied to understanding the control by landscape fire activity of pyrophobic tropical rain forest and pyrophytic eucalypt savanna boundaries, which are often separated by tall eucalypt forests. We evaluate the microclimate of three vegetation types across an elevational gradient and their relative fire risk as measured by McArthur's Forest Fire Danger Index (FFDI). Microclimatic data were collected from rain forest, tall eucalypt forest and savanna sites on eight vegetation boundaries throughout the humid tropics in north Queensland over a 3‐year period and were compared with data from a nearby meteorological station. There was a clear annual pattern in daily FFDI with highest values in the austral winter dry season and lowest values in the austral summer wet season. There was a strong association of the meteorological station FFDI values with those from the three vegetation types, albeit they were substantially lower. The rank order of FFDI values among the vegetation types decreased from savanna, tall eucalypt forest, then rain forest, a pattern that was consistent across each transect. Only very rarely would rain forest be flammable, despite being adjacent to highly flammable savannas. These results demonstrate the very strong effect of vegetation type on microclimate and fire risk, compared with the weak effect of elevation, consistent with a fire–vegetation feedback. This study is the first demonstration of how vegetation type influences microclimate and fire risk across a topographically complex tropical forest–savanna gradient.     

Effect of fire regime on plant abundance in a tropical eucalypt savanna of north‐eastern Australia

Abstract Changes in plant abundance within a eucalypt savanna of north‐eastern Australia were studied using a manipulative fire experiment. Three fire regimes were compared between 1997 and 2001: (i) control, savanna burnt in the mid‐dry season (July) 1997 only; (ii) early burnt, savanna burnt in the mid‐dry season 1997 and early dry season (May) 1999; and (iii) late burnt, savanna burnt in the mid‐dry season 1997 and late dry season (October) 1999. Five annual surveys of permanent plots detected stability in the abundance of most species, irrespective of fire regime. However, a significant increase in the abundance of several subshrubs, ephemeral and twining perennial forbs, and grasses occurred in the first year after fire, particularly after late dry season fires. The abundance of these species declined toward prefire levels in the second year after fire. The dominant grass Heteropogon triticeus significantly declined in abundance with fire intervals of 4 years. The density of trees (>2 m tall) significantly increased in the absence of fire for 4 years, because of the growth of saplings; and the basal area of the dominant tree Corymbia clarksoniana significantly increased over the 5‐year study, irrespective of fire regime. Conservation management of these savannas will need to balance the role of regular fires in maintaining the diversity of herbaceous species with the requirement of fire intervals of at least 4‐years for allowing the growth of saplings >2 m in height. Whereas late dry season fires may cause some tree mortality, the use of occasional late fires may help maintain sustainable populations of many grasses and forbs.     

Hidden crown jewels: the role of tree crowns for bryophyte and lichen species richness in sycamore maple wooded pastures

Tree crowns typically cover the vast majority of the surface area of trees, but they are rarely considered in diversity surveys of epiphytic bryophytes and lichens, especially in temperate Europe. Usually only stems are sampled. We assessed the number of bryophyte and lichen species on stems and in crowns of 80 solitary sycamore maple trees (Acer pseudoplatanus) at six sites in wooded pastures in the northern Alps. The total number of species detected per tree ranged from 13 to 60 for bryophytes, from 25 to 67 for lichens, and from 42 to 104 for bryophytes and lichens considered together. At the tree level, 29 % of bryophyte and 61 % of lichen species were recorded only in the crown. Considering all sampled trees together, only 4 % of bryophyte, compared to 34 % of lichen species, were never recorded on the stem. Five out of 10 red-listed bryophyte species and 29 out of 39 red-listed lichen species were more frequent in crowns. The species richness detected per tree was unexpectedly high, whereas the proportion of exclusive crown species was similar to studies from forest trees. For bryophytes, in contrast to lichens, sampling several stems can give a good estimation of the species present at a site. However, frequency estimates may be highly biased for lichens and bryophytes if crowns are not considered. Our study demonstrates that tree crowns need to be considered in research on these taxa, especially in biodiversity surveys and in conservation tasks involving lichens and to a lesser degree also bryophytes.