Epiphytic Lichen Richness Research Articles

Conservation of unmanaged pan-European forest landscapes as a priority natural heritage for epiphytic lichens at different ecological and biogeographical scales - A review

The European forestry landscape represents an important priority for biodiversity and needs adequate management strategies. The main focus of this review is the importance of European forest areas for epiphytic lichen richness under the impact of different management practices assessed in the biogeographical and ecological regions of Europe. In total, 88 scientific articles were reviewed (based on the first author’s archive and by query in Web of Science), which examined epiphytic lichen richness within managed and unmanaged European forests. Biogeographical and ecological regions of Europe were not taken into account in the reviewed articles, but we used them for statistical analyses in the present work according to the geographical position of the investigated sites published in the reviewed articles. We also analysed the dissimilarities in epiphytic lichen richness among European biogeographical and ecological regions. Additionally, we analysed the impact of different silvicultural management practices (within assessed forests) on epiphytic lichen richness across the different biogeographical and ecological regions of Europe. The main results indicate that epiphytic lichen richness is significantly different across the biogeographical and ecological regions of Europe. Epiphytic lichen richness is significantly greater in Western European broadleaf forests in the Carpathian and Caledonian mountain areas and significantly lower in the Central European mixed forests and East European forest steppe. Management practices applied within studied forests had a negative impact on epiphytic lichen richness, while epiphytic lichen richness was higher within unmanaged forests. The main conclusion is that forests within highland areas of Europe that are not subjected to anthropogenic activities or management practices harbour greater epiphytic lichen richness and therefore should be conserved and protected due to their biological and ecological importance.

Potential implications of shortened rotation length for forest birds, bryophytes, lichens and vascular plants: An example from southern Swedish production forests

The rotation lengths of intensively managed production forests may be altered to achieve a variety of goals, with correspondingly implications for biodiversity. Here we consider the potential implications of shortened rotation times for biodiversity in planted monocultures of the two most common production tree species in Sweden, Scots pine (Pinus sylvestris) and Norway spruce (Picea abies). To do so we surveyed bird, bryophyte, epiphytic lichen and vascular plant diversity in 80 and 55-year-old stands; stand ages which approximate present-day and potential future rotation lengths in this region respectively. We found clear differences in the species communities of the 55 compared to the 80-year-old stands for both understory species and epiphytes, but not for birds. Nevertheless, bird species richness was still highest in the 80-year-old Norway spruce dominated stands. Dead wood amount was also highest the 80-year-old Norway spruce stands. Highest species richness of epiphytic lichens was found in 80-year-old Scots pine stands. However, 55-year-old Scots pine stands had a higher understory species richness and diversity than the older Scots pine stands, including a larger number of open land species. The 80-year-old forest stands examined may be considered old with respect to production forest rotation lengths in Sweden but are relatively young when comparing stand ages of unmanaged natural forest stands. Nevertheless, our results indicate that shortening the rotation time of Scots pine and Norway spruce, in this part of Sweden from 80 to 55 years, could have important consequences for forest biodiversity. These consequences are primarily inferred from the likely implications from shortened rotations for lichens community composition and diversity in both Norway spruce and Scots pine stands, as well as impacts on understory plant species in Norway spruce stands.

Recovery kinetics of epiphytic lichen diversity after dieback during a continuously wet season

Epiphytic lichens are considered sensitive indicators of environmental change. Excess water is known to depress their photosynthesis, but the effect of long-lasting rain on species richness of epiphytic lichens is rarely reported. By annually repeated records of macrolichen species richness on tree trunks over a period of 33 years that included one long rainy season in year 2000, a strong decline in macrolichen richness on tree trunks was detected after the unusually wet autumn. Afterwards, the lichen richness slowly recovered, but had not yet fully recovered 19 years after the dieback. Thereby, long rainy periods can cause lasting depression in epiphytic lichen richness, and continuous rain should be considered a possible threat to lichens in regions like northern Europe where global change predicts enhanced rainfall frequency.

What is good for birds is not always good for lichens: Interactions between forest structure and species richness in managed boreal forests

Forestry in the boreal region increasingly replaces natural disturbances in shaping biodiversity. Large-scale removal of small diameter trees (thinning), is ubiquitous in northern European forestry, yet an understanding of how it relates to biodiversity across taxa is lacking. To address this, we examined how two forest structural elements, commonly impacted by forest thinning (i.e. vegetation density in the understory and overstory), are correlated with the species richness of forest-dwelling birds and epiphytic lichens. These taxa were chosen because they likely have opposing habitat demands: epiphytic lichens potentially benefitting from more light in open forests, and birds benefitting from structurally complex forests for foraging, nesting and cover. We used remote sensing tools, already applied in forestry planning, to see if management recommendations could be found that balance the needs of both taxonomic groups within the same forests. Our results show that richness in epiphytic lichens and forest-dwelling birds is not correlated and that a high species richness (~15 birds & ~40 epiphytic lichens) in both taxa is only predicted in the same stand under a specific vertical distribution and density of the forest’s vegetation: this occurs when the vegetation above 3 m is open (to promote epiphytic lichen richness) and below 3 m is dense (to promote bird richness). Dense vegetation up to 7 m above ground has a diverging effect on forest birds (positive) and epiphytic lichens (negative). A larger stem diameter further increases species richness in both taxa. Our study provides results that are directly implementable in forestry planning over large areas with the help of remote sensing tools (LiDAR).

Factors Influencing Epiphytic Lichen Species Distribution in a Managed Mediterranean Pinus nigra Arnold Forest

Lichens have important ecological functions in black pine forests, such as nitrogen fixation and nutrient cycling. Understanding lichen diversity could provide a better understanding of black pine ecosystems. The aim of this study was to identify the factors affecting the composition of lichen communities and their specific diversity in Mediterranean black pine forests. Research was conducted in 48 sampling plots. For the analysis, presence–absence and frequency data of lichen species were used. For stand level analysis, four community composition tables were created. We used bioclimate, topography, stand, and parent rock as variables. A total of 33 epiphytic lichen species were identified in the black pine forests from 282 sampled trees. Indicator lichen species were determined according to geographic region and stand age classes. Hypocenomyce scalaris was found to be an indicator species for old forests. Frequency data were more useful for revealing lichen species composition than presence–absence data. Of the topographic variables, elevation was the most prominent and had the highest explanation ratio for the composition of lichen species with a coefficient of correlation (R2) value of 0.49. Significantly positive (p < 0.001) relationships were found between epiphytic lichen richness and tree crown height, tree height, and bark pH. Our results revealed that to retain the trees in the stands rich in lichen species diversity is recommended in the managed forests.

Non-dominant trees significantly enhance species richness of epiphytic lichens in subtropical forests of southwest China

Abstract Host species has an important influence on the distribution of epiphytic lichens in forests. However, the importance of non-dominant trees in shaping lichen communities has been poorly studied owing to the relative rarity of individuals. The importance of dominant and non-dominant trees for distribution of epiphytic lichens was determined in eight subtropical forests in southwestern China. Dominant trees supported more abundant total and exclusive lichen species only in secondary forests. The occurrence of non-dominant trees promoted lichen diversity within forest types and influenced lichen communities on both tree groups. The effects of total tree species on lichen distribution largely resulted from the presence of non-dominant trees. Dominant and non-dominant trees supported distinct lichen assemblages within forest types, and ordination analyses showed a clear separation. Our study, therefore, reinforces the importance of non-dominant trees for conserving epiphytic lichens, and highlights that lichen assemblages are shaped by both dominant and non-dominant trees.

Small, fragmented native oak forests have better preserved epiphytic lichen communities than tree plantations in a temperate sub-oceanic Mediterranean climate region

Abstract In increasingly fragmented landscapes tree plantations are thought to help maintain forest continuity among native patches. Epiphytic lichens are one of the most sensitive groups to change in forest conditions, but the effects of this management practice are still poorly studied in temperate regions. In this study we compared epiphytic lichen diversity among small patches of old, native oaks (Quercus pyrenaica) and young, planted chestnuts (Castanea sativa) and Monterey pines (Pinus radiata) in a major forestry area in northwestern Spain. Our results showed that the richness of epiphytic lichens was higher in C. sativa than Q. pyrenaica, but that the latter had more lichen species typical of mature forests, including macrocyanolichens. Overall, C. sativa plantations had more species typical of well-lit, dry environments than Q. pyrenaica or P. radiata, which is likely caused by the natural architecture of each tree species, and to differences in forest age and management. At the tree level, Q. py...

The impact of pollutants from a major northern highway on an adjacent hardwood forest.

Emissions of pollutants from highways can exert multiple stresses on adjacent ecosystems. In this study air concentrations of NO2 and throughfall deposition of inorganic N (NO3- and NH4+), SO42-, Cl-, base cations and several metals were measured in all four seasons along a 1.5km hardwood forest gradient extending away from a major highway (Highway 401) in southern Ontario, Canada. Soil and lichen chemistry and herbaceous plant and epiphytic lichen species composition were measured within the hardwood forest to evaluate impacts of these pollutants. Air concentrations of NO2 and deposition of inorganic N, Cl-, base cations and Cu and Zn in throughfall were significantly elevated within 100m of the road compared with the more distant sites. Concentrations of several pollutants including N (and δ15N), Na+, Al and Fe in epiphytic lichen tissue decreased with distance from the highway, and epiphytic lichen richness was lower at sites within 100m of the road. Despite high throughfall inputs of >15kgNha-1y-1 and 100kgNa+ha-1y-1 within 33m of the highway, for example, there was no significant difference in soil chemistry amongst sites. Plant community composition at sites within 80m of the highway differed from sites located further from the road, but it is unclear whether differences were due to highway emissions or were a result of natural forest edge effects.

A survey method for assessing the richness of epiphytic lichens using growth forms

The species richness of epiphytic lichens is continuously decreasing by degradation and loss of habitat. Considering that taxonomic identification of all species is time and resource consuming, rapid assessment methods to extrapolate the total number of species are needed for practical conservation. This paper describes an alternative method using the correlation between lichens growth forms and species richness. The study was conducted in 406 forest stands located in Central Spain, covering a wide range of mediterranean-climate ecosystem regions, management intensity levels, canopy cover conditions, and tree sizes. The presence/absence of epiphytic lichens was determined in 6090 trees, which were dominated by oak species (Quercus ilex, Q. faginea, and Q. pyrenaica). In all type of forests, the diversity of growth forms was positively correlated with the total epiphytic lichen richness. In all cases, species richness increased in non-managed forest stands with dense canopies. Thus, we propose the use of lichen growth forms as a helpful surrogate of species richness to detect potentially conservation priority areas in the Mediterranean region.

The value of retained Scots pines and their dead wood legacies for lichen diversity in clear-cut forests: The effects of retention level and prescribed burning

To mitigate the negative impact of intensive forest management on biodiversity, several silvicultural measures are used in production forests. We studied the effect of two such treatments, green-tree retention and prescribed burning on epiphytic lichens 11years after timber harvests in Scots pine (Pinus sylvestris) dominated forests of middle boreal Finland. Lichen assemblages were sampled from retained Scots pines and their dead wood legacies (snags and logs) below a height of 2m. Twelve study sites were harvested with two retention levels (10 and 50m3/ha) and six uncut old-growth sites were used as controls. Prescribed burning was applied to half of the sites immediately after harvest. A total of 42 and 65 lichen species was found in the burned (B) and unburned (UB) sites with 10m3/ha retention, 41 and 74 in the B and UB sites with 50m3/ha retention, and 63 and 80 in the B and UB controls respectively. Mean species richness per tree did not differ between the control and harvested sites at either retention level on any of the substratum types. However, microlichen species richness was significantly lower on harvested sites on both living trees and snags. The species richness of both micro- and macrolichens was lower on burned sites. In addition, the composition of lichen assemblages differed between burned and unburned sites. The negative effect of fire was significant for lichens growing on living trees and snags, but not on logs. We conclude that the retention of Scots pines is a successful measure for maintaining epiphytic lichen richness over a timescale of 11years after harvest, and that roughly equal numbers of living trees and their dead wood legacies are needed to support the lichen species richness on harvested sites. Given that the negative effect of prescribed burning is relatively longer for lichens than for other wood-dependent organisms, burning should not be applied to stands with particularly rich lichen communities. However, these conclusions may have limitations as our inventory was limited to the lowermost part of tree trunks only. Hence inventories of whole trees are useful to assess the full effect of prescribed burning. Our results also imply that the specific impact of prescribed burning for specialist lichen species should be evaluated over a longer timescale.

Relationships between the diversity patterns of vascular plants, lichens and invertebrates in the Central Asian forest-steppe ecotone

The Central Asian forest-steppe ecotone has been exposed to large alterations in grazing pressure in the last two decades, but the consequences for biodiversity have not been studied so far. We analyzed the biodiversity of the edges and the interior of Siberian larch forests in the forest-steppes of eastern Kazakhstan (Saur, Kazakh Altai) and western Mongolia (Mongolian Altai, Khangai) across different groups of organisms (vascular plants, epiphytic lichens, soil macroarthropods, oribatid mites, moths). The species richness of these groups was related to each other only at the forest edge, but not in the interior. Species richness of vascular plants, soil macroarthropods and oribatid mites at the forest edges was positively correlated. This indicates that these ground-inhabiting groups of organisms responded similarly to the variation in the grazing pressure of livestock, which is kept at spatially varying densities by mostly nomadic or transhumant herders. The species richness of epiphytic lichens was only positively correlated with that of vascular plants, and the richness of the (volant) moths was not correlated with that of any other group. The complete lack of correlation between the diversity of groups of organisms in the forest interior suggests that the diversity of the five studied groups is controlled by specific environmental factors, including light and moisture. Except for the Mongolian Altai, which was subjected to the highest grazing pressure, vascular plants, lichens, soil macroarthropods, and moths had a higher diversity at the edges than in the interior; the opposite was true for the oribatid mites. The latter probably benefit from the higher soil moisture inside the forest, whereas the other four groups are favored by increased availability of light, the proximity to the steppe with a partial mixing of species pools, and the soil macroarthropods also by increased dung abundance.

Estimating epiphytic lichen richness by single families in Mediterranean forests

The current trend of declining epiphytic richness caused by human activities (forest fragmentation, logging, agriculture, and livestock grazing) and the greater efforts required to sample and identify the most inconspicuous species have necessitated the use of indicators of the species richness. In this study, we examined the potential of predicting epiphytic lichen richness based on the richness of a single taxon (family) of the most conspicuous lichens (macrolichens) in Mediterranean woodlands. Since our working hypothesis is that the richness of some conspicuous elements is tightly connected with the total richness, we expect this connection is maintained even after composition shifts (for instance composition changes between coniferous and oak forests). In order to control the large set of confounding factors at macro- and microclimate scales our present study was conducted in 504 forest stands, which represented a wide range of Mediterranean climates, management intensity levels, canopy cover types, and tree sizes. The presence/absence of epiphytic lichens were determined in 7560 trees, which were dominated by coniferous (Pinus nigra and P. sylvestris) and oak (Quercus ilex ssp. ballota, Q. faginea, and Q. pyrenaica) species. In oak forests, the increased richness of Collemataceae and the complex known as “rest of Peltigerales” was followed by an increase in the overall epiphytic richness, whereas there was a strong positive correlation between Parmeliaceae and total epiphytic richness in coniferous forests. In both cases, the richness of these predictors increased in well-preserved forest stands with dense canopies. Thus, we propose the potential use of Parmeliaceae (for coniferous forests) and the Collemataceae and the “rest of Peltigerales” (for oak forests) as indicators in the Mediterranean region because they have a cosmopolitan distribution, grow in a wide range of environmental conditions, and are correlated with changes in the epiphytic richness caused by forest disturbances.

Species Richness of Epiphytic Lichens in Coniferous Forests: the Effect of Canopy Openness

Epiphytic lichen communities were studied in old coniferous forests across Estonia to find out the effect of site openness on the lichen species richness. All lichen species were recorded on the basal 2 m of 105 Picea abies and 105 Pinus sylvetsris trunks. The canopy openness readings were taken with a spherical densiometer around every sampled tree. The number of species on the trunks of both tree species increased significantly with increasing canopy openness. Nine lichen species, e.g. Cladonia cenotea, Lecidea nylanderi, Platismatia glauca, were significantly favoured by higher light availability, and only one species, Coenogonium pineti, by lower light availability. The results demonstrate that in the coniferous forests higher light availability supports higher species richness of epiphytic lichens on the lower trunk of trees.

Large-scale patterns of epiphytic lichen species richness: Photobiont-dependent response to climate and forest structure

Lichens are composite organisms consisting of a symbiotic association of a fungus with a photosynthetic partner. Although the photobiont type is a key life-history trait, tests of the potential differential role of the main photobiont types in shaping large-scale patterns of lichen species richness are still absent. The aim of the study was to test the influences of forest structure and climate on epiphytic lichen species richness across Italy and to see whether these relationships change for groups of species sharing different photobiont types. Regional species richness of epiphytic lichens divided into three main photobiont types (i.e. chlorococcoid green algae, cyanobacteria, and Trentepohlia algae) was retrieved for each of the 20 administrative regions. Multiple linear regression was used to quantify the effect of climate and forest structure, and their potential interaction, on the regional species richness for the three photobiont types, accounting also for the effect of regional area. Regional species richness was associated with both climate and forest structure variables but the relationships with both factors were largely photobiont dependent. Regional area and precipitation were the only predictors included in all the models, confirming the strong dependence of lichens on atmospheric water supply, irrespective of the photobiont type. Number of species with chlorococcoid green algae were further positively associated with cover of high forest, whilst lichens with Trentepohlia were further enhanced by warm temperatures. Cyanolichen species richness was only related to area and precipitation. Our study shed light on the relative importance of climate and forest structure on lichen species richness patterns at the macroscale, showing a differential response of the photobiont types to various environmental determinants. This differential response suggested that the current and future impacts of global change on lichens cannot be generalized and that species richness response will be likely dependent on the photobiont type.

Isolation affects tree-scale epiphytic lichen community structure on New Zealand mountain beech trees

Question: Is epiphytic lichen community structure significantly affected by isolation from source community? Location: Foothills of the Southern Alps, South Island, New Zealand. Methods: Epiphytic lichen richness and environmental variables were measured on 382, young Nothofagus solandri var. cliffortioides (Hook. f.) Poole (mountain beech) trees that had recently colonized grassland adjacent to a forest remnant. Richness and the presence of individual lichen taxa were modelled as a function of isolation from the forest fragment, tree size and other habitat conditions. Results: Richness of epiphytic lichen communities was negatively related to tree isolation, although this effect was much smaller than the effects of tree size and other local (tree-scale) habitat conditions. Different lichen taxa responded in different ways to isolation, area effects and local habitat conditions. Conclusions: This study shows that many epiphytic lichens on mountain beech are limited in their ability to colonize new substrate, even over distances of less than 1 km, which may be due to limitation in dispersal and/or establishment. Lichens are greatly influenced by local habitat conditions, such as tree size, and in this particular environment their negative interaction with sooty moulds is an important driver of community structure.

Lichen diversity and red-listed lichen species relationships with tree species and diameter in wooded meadows

This study provides a unique large dataset of total epiphytic lichen diversity (fruticose, foliose and crustose species) and composition on 1,294 trees of 17 tree species in wooded meadows in Sweden and Estonia, the Baltic region. The inventory (25,380 observations and 246 lichen taxa) clearly illustrated that Ulmus minor, Quercus robur and Fraxinus excelsior contributed most significantly to epiphytic lichen richness and number of red-listed species. In Sweden, average single tree α richness was 22.2 on Ulmus (only in Sweden), 21.6 on Quercus (25.0 in Estonia) and 19.8 on Fraxinus (16.7 in Estonia), respectively. Ulmus hosted on average one red-listed species per tree, compared with 0.7 on Fraxinus (0.6 in Estonia), 0.4 on Quercus (0.7 in Estonia) and only 0.05 on Betula (same in Estonia). Lichen species composition and the average number of red-listed lichens were influenced by tree diameter on Fraxinus and Quercus, whilst no such pattern was evident on Ulmus. Randomized species accumulation curves of the dominating tree species illustrated that Fraxinus, Quercus and Ulmus supported α dominated lichen communities where individual trees hosted a substantial part of the total richness. Betula, on the other hand, supported β dominated communities where individual trees tended to be dissimilar and, therefore, more of the total richness existed as species turnover among host trees. Lichen species composition was influenced by tree species, and most notably, lichen species on Ulmus had a strong consistent clumping in ordination graphs, with many rare and red-listed lichens. The broadleaved deciduous trees within the wooded meadows clearly contribute greatly to the biodiversity of the Baltic region.

Factors controlling epiphytic lichen biomass during postfire succession in black spruce boreal forests

Alectorioid lichens are the dominant group of epiphytic lichens in boreal forests. Epiphytic lichen richness and abundance generally increase with stand age and within-stand heterogeneity. The objective of this study was to evaluate the importance of time elapsed since the last fire, stand structure, tree size, tree age, and branch height for epiphytic lichen biomass of the boreal forest of western Quebec. We sampled 12 sites belonging to four forest age classes (from 50 to >200 years). We assessed epiphytic lichen biomass of three species groups ( Bryoria , Evernia , and Usnea ) on 12 trees in each site. Our results showed that biomass of Bryoria and Usnea was higher in intermediate stages (between 101 and 200 years) compared with younger (50–100 years) and older (>200 years) stages. Biomass of the three species groups was greater on larger diameter trees (>16 cm) compared with smaller ones (<16 cm). These results indicate that the protection of postfire stands aged between 101 and 200 year should be prioritized to maintain the functional role of epiphytic lichens in managed landscapes.

Richness of epiphytic lichens in differently agedPicea abiesplantations situated in the oceanic region of Central Norway

AbstractThis study aims to investigate patterns of species richness and abundance in relation to stand age in 71Picea abiesplantations, aged between 9 and 85 years, situated in the oceanic region of Central Norway. The study has shown that plantations within the oceanic spruce forests can support a relatively high number of epiphytic lichen species. Some of the oldest plantations hosted several old-forest associated species, e.g.Hypogymnia vittata, Lobaria pulmonaria, Pseudocyphellaria crocataandRamalina thrausta. The number of species was influenced significantly by stand age and increased rapidly in stands <20 years old. Stands >30 years old showed no clear increase in species number, except for a high number of species in the two oldest stands. The colonization pattern could be characterized as an additional entrance of species, rather than by a replacement sequence. The probability of occurrence increased steeply at young stand ages (<20 years) forBryoriaspp.,Cavernularia hultenii, Platismatia glauca, Parmelia sulcataandUsneaspp. A lower rate of colonization was characteristic forAlectoria sarmentosa, Parmelia saxatilisandPlatismatia norvegica. The cover of foliose lichens on the branches showed an almost unimodal response to stand age. The cover of lichens was highest on branches in middle-aged plantations. The reason for the lower lichen cover in late successional stages, compared to middle-aged stands, could be due to reduced light in the lower canopy of mature plantations. Increased rotation cycle, creation of gaps and short distance to sources of propagules are factors suggested to promote species richness and abundance in forest plantations.

Tree and stand level variables influencing diversity of lichens on temperate broad-leaved trees in boreo-nemoral floodplain forests

Tree and stand level variables affecting the species richness, cover and composition of epiphytic lichens on temperate broad-leaved trees (Fraxinus excelsior, Quercus robur, Tilia cordata, Ulmus glabra, and U. laevis) were analysed in floodplain forest stands in Estonia. The effect of tree species, substrate characteristics, and stand and regional variables were tested by partial canonical correspondence analysis (pCCA) and by general linear mixed models (GLMM). The most pronounced factors affecting the species richness, cover and composition of epiphytic lichens are acidity of tree bark, bryophyte cover and circumference of tree stems. Stand level characteristics have less effects on the species richness of epiphytic lichens, however, lichen cover and composition was influenced by stand age and light availability. The boreo-nemoral floodplain forests represent valuable habitats for epiphytic lichens. As substrate-related factors influence the species diversity of lichens on temperate broad-leaved trees differently, it is important to consider the effect of each tree species in biodiversity and conservation studies of lichens.

Disturbance effects on diversity of epiphytes and moths in a montane forest in Ecuador

We sampled the diversity of epiphytes (lichens, bryophytes, vascular plants) and moths (Geometridae, Arctiidae) in mature and recovering forest and in open vegetation in the montane belt in Ecuador. No uniform pattern of change in species richness was detected among the different taxonomic groups with increasing disturbance. Species richness of epiphytic bryophytes and vascular plants declined significantly from mature forest towards open vegetation. In contrast, species richness of epiphytic lichens did not change with increasing forest alteration, while that of geometrid moths was significantly higher in recovering forest compared with mature forest and open habitats. Arctiidae were significantly more species-rich in recovering forest and open vegetation than mature forest. Hence, for some organisms, modified habitats may play an important role for biodiversity conservation in the Andes, whereas others suffer from habitat disturbance. However, trends of changes in species composition following deforestation were surprisingly concordant across most studied epiphyte and moth taxa.