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.     

Forest edge-interior differentiation in the epiphytic lichen diversity of the forest steppe in the Khangai Mountains,Mongolia

Aims The effects of traditional land use by mobile livestock keepers on biodiversity in forest steppe ecotones are insufficiently studied. Epiphytes are an important part of forest plant diversity. Here we analyze differences in the diversity and composition of the epiphytic lichen vegetation between the edge and the interior of Siberian larch forests in the Khangai Mountains, western Mongolia, which are highly subdivided into patches. We asked whether the epiphytic lichen vegetation at the forest edge differs significantly from that in the interior, whether the edge is inhabited by more nitrophilous species than the interior and whether the density of nomad camps around the forest affects epiphytic lichen diversity.Methods Cover percentages of epiphytic lichen species were recorded from 20 trees per plot on 6 plots in the interior and 6 plots at the edge of Larix sibirica forests. The position of nomad summer camps was surveyed using Global Positioning System. Data were analyzed with pairwise significance tests, analysis of similarities, nonmetric multidimensional scaling and canonical correspondence analysis.Important findings The composition of the epiphytic lichen vegetation clearly differed between the two habitats, with more species being more frequent at the edge than in the interior. However, there was no difference in species richness (α-diversity). The epiphyte vegetation at the edge was more uniform and characterized by lower variation of tree-level α-diversity and lower β-diversity than in the interior. At the edge, only nitrophytic lichens were dominant, whereas in the interior, nitrophytes and acidophytes were among the dominant species. This pattern is probably attributable to the spatial heterogeneity of the intensity of forest grazing and was shown to be influenced by the density of nomad summer camps in the vicinity of the forests. Tree-level α-diversity increased with stem diameter, but high-diameter trees were rare. The results suggest that the present level of forest patchiness and the effect of forest grazing increases the diversity of epiphytic lichens on the landscape level, while logging of high-diameter trees reduces lichen diversity.     

Lichens and bryophytes as indicators of old‐growth features in Mediterranean forests

Abstract

This study is focused on the selection of variables affecting lichen and bryophyte diversity in Mediterranean deciduous forests. Plots representing two forest types (Fagus sylvatica and Quercus cerris forests) and two forest continuity categories (old‐growth (OG) and non‐OG forests) were selected in the Cilento and Vallo di Diano National Park (Italy). The presence and the abundance of bryophytes and epiphytic lichens were recorded. Structural variables of the forests and vascular plant species richness have been used as predictors. A strong positive correspondence between the two groups of organisms was found. Higher species richness and the distribution of rare species are related to OG stands, while a qualitative (species composition) rather than a quantitative (species richness) difference between the two forest types was observed. Some species elsewhere considered as indicators of forest continuity, such as Lobaria pulmonaria, Antitrichia curtipendula, and Homalothecium sericeum, are associated with OG forests, independently from forest type, suggesting that they can be regarded as suitable indicators also in Mediterranean forests. Finally, our results suggest that old trees, high levels of basal area, a broad range of diameter classes, and high understory diversity are the main structural features affecting cryptogamic communities, while no correlation was found with the occurrence of deadwood.     

Aspen succession and nitrogen loading: a case for epiphytic lichens as bioindicators in the Rocky Mountains,USA

Question: Can lichen communities be used to assess short‐ and long‐term factors affecting seral quaking aspen (Populus tremuloides) communities at the landscape scale? Location: Bear River Range, within the Rocky Mountains, in northern Utah and southern Idaho, USA. Method: Forty‐seven randomly selected mid‐elevation aspen stands were sampled for lichens and stand conditions. Plots were characterized according to tree species cover, basal area, stand age, bole scarring, tree damage, and presence of lichen species. We also recorded ammonia emissions with passive sensors at 25 urban and agricultural sites throughout an adjacent populated valley upwind of the forest stands. Nonmetric multidimensional scaling (NMS) ordination was used to evaluate an array of 20 variables suspected to influence lichen communities. Results: In NMS, forest succession explained most variance in lichen composition and abundance, although atmospheric nitrogen from local agricultural and urban sources also significantly influenced the lichen communities. Abundance of nitrophilous lichen species decreased with distance from peak ammonia sources and the urban center in all aspen succession classes. One lichen, Phaeophyscia nigricans, was found to be an effective bioindicator of nitrogen loading. Conclusions: Lichen communities in this landscape assessment of aspen forests showed clear responses to long‐term (stand succession) and short‐term (nitrogen deposition) influences. At the same time, several environmental factors (e.g. tree damage and scarring, distance to valley, topography, and stand age) had little influence on these same lichen communities. We recommend further use of epiphytic lichens as bioindicators of dynamic forest conditions.     

Effects of forest management on epiphytic lichen diversity in Mediterranean forests

Question: What are the responses of epiphytic lichens to the intensity of management along a large environmental gradient in Mediterranean Quercus forests? Location: Central Spain. Methods: This study was carried out on 4590 trees located in 306 forest stands dominated by Quercus faginea or Quercus ilex ssp. ballota. The effect of forest management and other predictor variables on several species diversity indicators were studied. Variables modelled were total species richness, cyanolichen richness and community composition. A large number of predictor variables were included: forest fragmentation (patch size, stand variability), climate and topographic (altitude, slope, sun radiation, annual rainfall and mean annual temperature) and intensity of management. General linear models and constrained ordination techniques were used to model community traits and species composition, respectively. Results: Total richness and especially cyanolichens richness were significantly and negatively affected by the intensity of management. Lichen composition was influenced by management intensity, climatic and topographic variables and stand variability. Conclusions: In Mediterranean forests, human activities related to forestry, agricultural and livestock use cause impoverishment of lichen communities, including the local disappearance of the most demanding species. The conservation of unmanaged forests with a dense canopy is crucial for lichen diversity.     

Assessing the effects of forest management on epiphytic lichens in coppiced forests using different indicators

Abstract

Epiphytic lichens are one of the taxonomic groups most sensitive to forest management. Nevertheless, they have not yet been exhaustively included in the assessment of Sustainable Forest Management. This work aimed at evaluating the effects of forest management on epiphytic lichens in coppiced forests, exploring the spatial patterns of diversity and the composition of communities. Moreover, the goal was to compare the performance of four potential indicators for monitoring the effects of forest management on epiphytic lichens: total lichen diversity, species associated with intensive management, species associated with aged coppiced woodlands and Indicator Species Ratio (ISR). In humid Mediterranean Liguria, 50 sampling units were chosen in Castanea sativa and deciduous Quercus spp. forests subjected to different forest management practices: intensively managed coppice and aged coppice/high forest. The effect of forest management was evident in terms of species composition, since it was possible to find significantly associated species for each of the two management types. At each sampling site, the four indicators were calculated using Indicator Value Analysis and compared through correspondence analysis. The ISR was shown to be a more effective indicator, being independent of floristic composition and the occurrence of rare species.     

Edge and land-use effects on epiphytic lichen diversity in the forest-steppe ecotone of the Mongolian Altai

Epiphytic lichen diversity was studied in forests of Siberian larch (Larix sibirica) in the forest-steppe ecotone of the Mongolian Altai. These forests are utilized for livestock grazing, fuelwood collection and occasional logging by pastoral nomads. The density of nomad households in the proximity of the forests influences epiphytic lichen diversity more strongly than the position of sample trees in the forest interior or at the forest line to the steppe. This suggests that land use exerts a stronger effect on lichen diversity than the distinct gradient in microclimate between the forest interior and the forest edge. The co-occurrence of nitrophytes with anitrophytic acidophytes on a small spatial scale as well as higher N and Ca concentrations and pH values in the bark of larch trees at the forest edge than in the forest interior indicate that moderate livestock grazing increases the epiphytic lichen diversity due to an increase of the diversity of chemically different microhabitats. Preference of many lichen species (and of rare species in particular) for overmature and decaying trees suggests that logging and fuelwood collection has adverse effects on epiphytic lichen diversity. This adverse effect is likely to be more crucial for lichen diversity than the putatively positive effect of livestock grazing, since more lichens with a preference for old and decaying trees than for nitrogen-enriched bark were found. The present study is the first one investigating the impact of pastoral nomadism in Central Asia on epiphyte diversity.     

Diversity and succession of epiphytic macrolichen communities in low‐elevation managed conifer forests in Western Oregon

Abstract. We examined epiphytic macrolichen communities in Pseudotsuga menziesii (Douglas‐fir) forests across the western Oregon landscape for relationships to environmental gradients, stand age and structure, and commercial thinning. We used a retrospective, blocked design through the Coast and the western Cascade ranges of Oregon. Each of our 17 blocks consisted of a young, unthinned stand (age 50–110 yr); an adjacent, thinned stand of equivalent age; and an old‐growth stand (age > 200 yr). We found 110 epiphytic macrolichen taxa in the stands. Forage‐providing alectorioid lichens and the nitrogen‐fixing cyanolichen Lobaria oregana associated strongly with old‐growth stands and remnant old trees in younger stands (unthinned + thinned). Relative to unthinned stands, thinned stands had a slightly higher abundance of alectorioid lichens and a greater presence of Hypogymnia imshaugii. However, thinned stands hosted a lower landscape‐level (γ) diversity, lacking many species that occurred infrequently in the unthinned stands. Patterns in the lichen community composition correlated strongly with climatic gradients; the greatest variation in composition was between the Coast and Cascade ranges. The difference in communities between mountain ranges was greatest among stands 70–110 yr old, suggesting a difference in lichen successional dynamics between the ranges.     

亚热带常绿阔叶林附生地衣凋落物的物种多样性和生物量的边缘效应

地衣是亚热带山地森林系统附生生物类群的重要组成部分之一,对环境变化极其敏感。为了更好地了解附生地衣对森林边缘效应的响应,我们在云南哀牢山地区原生山地常绿阔叶林中,分别在距林缘5m、25m、50m和 100m处设立样地,收集附生大型地衣的凋落物1年;分析附生地衣凋落物的物种多样性和生物量、功能群特征和组成结构对林缘深度变化的响应特征。研究结果显示,边缘效应能够显著提高林缘附生地衣群落的物种多样性和生物量;其发生的距离最深不超过25m。林缘-林内梯度上,不同地衣功能群的响应模式具有各自的特征。排序分析表明仅在5m样地与其他样地之间存在显著差异,指示种分析则发现仅5m样地具有指示种。哀牢山原生林中边缘效应促进林缘附生地衣生长和分布的现象,可能与当地高湿环境削减了地衣的高光损伤以及以叶状和灌状类群为主的地衣个体受到风力破坏的程度相对较低有关。     

Litterfall of epiphytic macrolichens in Nothofagus forests of northern Patagonia,Argentina: Relation to stand age and precipitation

Abstract: The objective of this study was to analyse how stand age and precipitation influence abundance and diversity of epiphytic macrolichens in southern beech Nothofagus forests, estimated by lichen litter sampling. Five sites of Nothofagus dombeyi (Mirbel) Oersted were selected in Nahuel Huapi National Park, Argentina. At each site, lichen fragments from the forest floor were collected at 12.5 m² plots in pairs of young and mature N. dombeyi forest. Additionally, two sites with multi‐aged subalpine Nothofagus pumilio (Poepp. et Endl.) Krasser forest were investigated in a similar manner. Average litterfall biomass per stand varied from less than 1 kg ha^?1 in a young low‐precipitation stand to a maximum of 20 kg ha^?1 in a mature high‐precipitation stand. In places with higher precipitation, litterfall biomass in N. dombeyi forest was considerably higher in old stands as compared with young ones. In places with less than 2000 mm of precipitation, differences in biomass were less pronounced. Old humid stands contained about twice as many taxa in the litter as old low‐precipitation stands and young stands in general. Mature stands in low‐precipitation sites only contained 17% of the litter biomass as compared with mature stands in high‐precipitation sites. Epiphytic lichen composition changed from predominating fruticose lichens (Usnea spp. and Protousnea spp.) in low‐precipitation stands to Pseudocyphellaria spp., Nephroma spp. and other foliose lichens, in the high‐precipitation stands. There were no clear differences in the proportion of fruticose and foliose lichens between young and old stands. Fruticose lichens dominated litter biomass in both N. pumilio sites.     

云南哀牢山原生林及次生林群落附生地衣物种多样性与分布

附生地衣是森林附生植物的重要类群之一, 在维护森林生态系统的物种多样性以及水分和养分循环等方面发挥着重要作用。作者于2005年12月至2006年5月利用树干取样法调查了云南哀牢山徐家坝地区原生山地常绿阔叶林及其次生群落栎类萌生林、滇山杨(Populus bonatii)林和花椒(Zanthoxylum bungeanum)人工林中525株不同种类和径级树木距地面 0–2.0 m处附生地衣的组成和分布, 并收集了各个群落地面上凋落的地衣, 分析了林冠层附生地衣的物种组成。研究结果表明, 该区森林中附生地衣物种比较丰富。共收集到附生地衣61种, 分属17科29属, 其中原生林、栎类萌生林、滇山杨林和花椒人工林分别有51、53、46和23种。在树干距地面 0–2.0 m位置, 各群落中的附生地衣组成明显不同;但在林冠层中, 各群落内的附生地衣基本相似。原生林中附生地衣种类较多, 但分布不均匀。树干附生地衣的Shannon-Wiener和Simpson多样性指数以栎类萌生林最高, 分别为2.71和0.89;花椒林和滇山杨林次之, 分别为2.43–2.45和0.88–0.89;原生林最低, 为1.25和0.67。树干方位、宿主种类和宿主径级等都对附生地衣的物种组成和多样性有着重要影响, 附生地衣更多地出现于树干南向方位, 云南越桔(Vaccinium duclouxii)的附生地衣最为丰富, 胸径5.0–25.0 cm的树木上附生地衣较多。哀牢山山地森林群落中丰富的附生地衣种类及物种多样性在维系本区山地森林生态系统生物多样性格局方面具有重要的作用。     

Bryophyte Species Diversity in Secondary Forests Dominated by the Introduced Species Spathodea campanulata Beauv. in Puerto Rico

The introduced tree species Spathodea campanulata (Bignoniaceae) forms novel forests in Puerto Rico, these having emerged after the abandonment of fields in the mid‐20th century and resulting in forests with a new species composition. We assessed bryophyte species richness in these novel forests and sought correlations with geological substrate, past land use, forest edge and patch area, forest structure, elevation, microhabitat diversity, tree species richness, and microclimatic conditions. Transects were established (edge and forest interior) in nine moist forest patches dominated by Spathodea in north‐central Puerto Rico. These Spathodea forest patches ranged from 0.6 to 9 ha. ANOVA, Chi‐square, correlation, and cluster analyses were used in data analyses. We found 57 bryophyte species. There was a significant difference in bryophyte richness among patches. Those on karst exhibited highest bryophyte richness due to microhabitat diversity, past land use, and shorter hydroperiods. Alluvial sites scored lowest in bryophyte species richness, and forest structure was important for bryophyte communities on these sites. Significant differences in temperature, relative humidity, and light intensity were observed between edge and forest interior. These appeared important for establishing bryophyte species cover but not richness and composition. Microhabitat diversity, patch area, and forest age were more related to bryophyte species richness than elevation, exposed edge, and tree species richness, regardless of geologic substrate. Collectively, Spathodea patches were similar to mature forests on the Island with respect to bryophyte species richness and composition. Novel Spathodea forests have conservation value due to their habitat suitability for bryophyte communities.     

Small logging roads do not restrict movements of forest rats in Bornean logged forests

Selective logging is driving the proliferation of roads throughout tropical rain forests, particularly narrow, unpaved logging roads. However, little is known about the extent of road edge effects or their influence on the movements of tropical understory animal species. Here, we used forest rats to address the following questions: (a) Does the occupancy of rats differ from road edges to forest interior within logged forests? (b) Do roads inhibit the movements of rats within these forests? We established trapping grids along a road edge‐to‐forest interior gradient at four roads and in three control sites within a logged forest in Sabah, Malaysia. To quantify the probability of road crossing, rats were captured, translocated across a road, and then recaptured on subsequent nights. We caught 216 individuals of eight species on 3,024 trap nights. Rat occupancy did not differ across the gradient from road edge to interior, and 48 percent of the 105 translocated individuals crossed the roads and were recaptured. This proportion was not significantly different from that of rats returning in control sites (38% of 60 individuals), suggesting that small roads were not barriers to rat movements within logged forests. Subadults were significantly more likely to return from translocation than adults in both road and control sites. Our results are encouraging for the ecology of small mammal communities in heavily logged forests, because small logging roads do not restrict the movements of rats and therefore are unlikely to create an edge effect or influence habitat selection.     

Epiphytic lichen synusiae and functional trait groups in boreo‐nemoral deciduous forests are influenced by host tree and environmental factors

Deciduous forests with temperate broad‐leaved tree species are particularily important in terms of biodiversity and its protection, but are threatened habitats in northern Europe. Using multivariate analyses we studied the effect of forest site type, environmental variables and host tree properties on epiphytic lichen synusiae as well as on the composition of species‐specific functional traits. Epiphytic lichens were examined on Acer platanoides, Fraxinus excelsior, Quercus robur, Tilia cordata, Ulmus glabra and U. laevis in two types of forests: Humulus‐type floodplain forests and Lunaria‐type boreo‐nemoral forests on the talus slopes of limestone escarpment (klint forests). Klint forests located near the seashore were under greater maritime influence compared to floodplain forests located in inland Estonia which experience stronger air temperature contrasts. In addition to stand level and climatic variables, tree level factors (bark pH, trunk circumference and cover of bryophytes) considerably affected the species composition of the lichen synusiae. Overall, 137 lichen species were recorded, including 14 red‐listed species characteristic of deciduous trees. We defined 13 lichen societies and showed their preference to forests of a specific site type and/or host tree properties. In forests of both types, most of the epiphytic lichens were crustose, and had apothecia as the fruit bodies and chlorococcoid algae as the photobiont. However, the proportion of lichens with a foliose or fruticose growth form, as well as the proportion of lichens with vegatative diaspores, were higher in floodplain forests. In klint forests with a stronger influence from the wind, crustose species completely dominated, while species with vegetative diaspores were rare and most species dispersed sexually. Lichens with Trentepohlia as the photobiont were characteristic of these forests, and lichens with lirellate ascomata were prevailing, indicating the great uniqueness of the kint forests for epiphytic lichens in the boreo‐nemoral region.     

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

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.     

The Mediterranean Quercus pyrenaica oak forest: a new habitat for the Capercaillie?

We described an extension of the known distribution range of the Cantabrian Capercaillie Tetrao urogallus cantabricus into an atypical area and habitat for the species. Nine Capercaillie leks and 14 cocks were registered in Mediterranean Quercus pyrenaica forests in an area of 1,500 km², of which 4,500 forest hectares were surveyed. At present, this population represents both the southern-most distribution for Capercaillie and the only one inhabiting Mediterranean Q. pyrenaica forests, what suggests a wider adaptation of this (sub)species than previously thought. This population and its habitat need to be better studied, as well as to be considered in conservation planning for Cantabrian Capercaillie.     

Stand characteristics and biodiversity indicators along the productivity gradient in boreal forests: Defining a critical set of indicators for the monitoring of habitat nature quality

Abstract

We quantified the effects of anthropogenic disturbances on the structure and biodiversity of boreal forests on acidic soils and created a statistically supported rational set of indicators to monitor the stand “naturalness”. For that, we surveyed various traits of tree layer, understory, herb layer, forest floor and several widely accepted biodiversity epiphytic indicators in 252 old‐aged boreal stands in Estonia, mostly dominated by Scots pine or Norway spruce. Multifactorial general linear model analyses showed that many forest characteristics and potential indicators were confounded by the gradient of soil productivity (reflected by the forest site type), local biogeographic gradients and also by stand age. Considering confounding effects, boreal forests in a near‐natural state have more large‐diameter trees (diameter at breast height >40 cm) and larger variety of diameter classes, higher proportion of spruce or deciduous trees, a larger amount of coarse woody debris in various stages, a more closed tree canopy and denser understory than managed mature forests. By increasing light availability above the field layer, forest management indirectly increases the coverage of herbs and lichens on the forest floor but reduces the alpha‐ and beta‐diversity of herbs and the proportion of graminoids. Human disturbances reduce the relative incidence of many commonly accepted biodiversity indicators such as indicator lichens, woodpeckers, wood‐dwelling insects or fungi on trees. The test for the predictive power of characteristics reacting on disturbance revealed that only a fraction of them appeared to be included in a diagnostic easy‐to‐apply set of indicators to assess the nature quality of boreal forest: the amount of dead wood, the proportion of deciduous trees, the presence of specially shaped trees and woodpeckers and, as an indicator of disturbances, the forest herb Melampyrum pratensis. Many of these indicators have already been implemented in practice.     

Climatic discrimination of Mediterranean broad-leaved sclerophyllous and deciduous forests in central Spain

Abstract. Climatic differences between three types of deciduous (Quercus pyrenaica) and three types of sclerophyllous (Quercus rotundifolia) Mediterranean forests in the Spanish Sistema Central were analyzed by means of Canonical Discriminant Analysis and Jancey's Discriminant Analysis, applied in successive steps to data from 252 meteorological stations. Climatic data included temperature and precipitation records as well as bioclimatic indices. Discriminant analysis was applied to broad-leaved sclerophyllous and deciduous forest communities sampled at each meteorological station using phytosociological methods. Annual and seasonal (summer, spring) water availability are the most important factor controlling the distribution of the two physiognomic forest types; southwestern associations of Quercus pyrenaica and Q. rotundifolia differ from their colder homologues by annual and monthly temperatures; western associations were separated from eastern ones in terms of annual and seasonal precipitation gradients. Discriminant analysis was a good technique to explore climatic gradients not shown by other general ordination or classification methods.     

Road edge effect on the abundance of the lizard <Emphasis Type="Italic">Gallotia galloti</Emphasis> (Sauria: Lacertidae) in two Canary Islands forests

Transportation infrastructure is a main cause of environmental change in forest landscapes worldwide. In the Canary Islands, a dense road system fragment the native Canarian pine and laurel forests causing potential changes in population densities of endemic lacertid lizards (genus Gallotia). Our aim was to assess road edge effects on relative abundance patterns of the endemic Gallotia galloti in both forests. We also explored the species–habitat relationships in this road-fragmentation context. We found that lizard relative density in relation to road edges differed between forests. Lizards were more abundant along edges and leeward interior, but virtually absent from the interior of the windward laurel forest. In the pine forest, lizards were present at three distances from edge, with a net decrease in abundance from edge to interior. These patterns may be explained partly by differences in vegetation structure regarding road proximity in each forest that potentially affect the helio- and thigmothermic character of G. galloti, and thus its habitat use. A general suggestion of this study is that road margins create corridors that may be used by native lizards for dispersal through inhospitable forest matrix. The high road density in Tenerife may have negative implications for the conservation of the genetic variability of G. galloti. At the island scale, increased communication between lizard populations through road corridors might increase homogenization of the gene pool. Ecological processes in which this lizard plays important roles may also experience changes along road edges.