Growth and vitality of epiphytic lichens

 We tested the hypothesis that changed microclimate at induced forest edges causes reduced growth of epiphytic lichens. Two foliose, green algal lichens were transplanted to the lower canopy of a mature Picea abies forest at six distances (2, 6.25, 12.5, 25, 50 and 100 m) from a clearcut. The biomass growth in Platismatia glauca (6.2% in 16 months) was 41% higher than in Lobaria pulmonaria (4.4%). We found no growth reduction near the forest edge. In contrast, the highest growth in both species occurred within 12 m from the edge. Further, fluorescence and chlorophyll measurements showed that lichen vitality was unaffected by distance from edge. The light intensity was 4.3 times higher at the edge than in the interior during the growing season, but there were only minor differences in air temperature and relative humidity. Monitoring of thallus water content revealed clear differences in both number and length of wetting and drying cycles. However, the total time with water content sufficient for photosynthetic activity was only slightly higher at the edge. The data thus indicate that our gradient in microclimate was too small to significantly affect lichen growth, and that lichens are largely metabolically inactive when large edge-interior contrasts in microclimate occur. Lichen response to forest edge microclimate results from intricate interactions among several biotic and abiotic factors. Linking data on lichen growth, microclimate and thallus water content with physiological measurements provides a framework for future studies of the mechanisms behind abiotic edge effects. Received: 15 April 1996 / Accepted: 21 June 1996     

Two-hundred-years old shoots of the epiphytic moss Brotherella henonii preserved in a bark pocket of the conifer Cryptomeria japonica

Abstract

Schistidium flaccidum (De Not.) Ochyra is recorded as new to Britain from Snowdon, Wales. It may be distinguished from the other British species of the genus by its poculiform capsule, absence of peristome and small, dense tufts. Notes are given on its separation from Grimmia anodon, which it resembles more closely.     

Abscisic acid concentrations and fluxes in droughted conifer saplings

We present the first study of abscisic acid (ABA) concentrations and fluxes in the xylem sap of conifers during a drought cycle. In both Pinus sylvestris and Picea sitchensis the concentration of ABA in the sap rose 11-fold as the drought progressed. There were clear diurnal trends in this concentration, which reached its maximum (6–8.ininol ABA m^?3) near the middle of the day. The fluxes of ABA were calculated by multiplying the xylem ABA concentration by the sap flow rate. The ABA fluxes in the droughted plants in the middle of the day were usually no higher than those of the controls, as a result of the very low sap flow in the droughted plants at that time. However, the ABA flux in the droughted plants was higher than in the controls in the morning, and we postulate that the stomata are responding to these ‘morning doses’ Stomatal conductance, g_s, could not be related statistically to leaf turgor or to the ABA flux. However, £s did display a negative exponential relationship with ABA concentration in the xylem. Pinus displayed more acclimation to drought than Picea, Its ABA concentration rose and its stomatal conductance fell at day 6 of the drought, as opposed to day 17 for Picea, and its osmotic potential fell during the drought treatment.     

Freezing avoidance mechanism of primordial shoots of conifer buds

Excised winter buds of very hardy fir supercooled to —30or — 35?C, though primordial shoots excised from thesewinter buds (freezing point: about —5.5?C) supercooledonly to —12 to — 14?C. Also, excised primordialshoots did not tolerate freezing, but were rather resistantto desiccation. Differential thermal analysis (DTA) of primordialshoots revealed that the capability of supercooling increasedwith decreasing water content and that no exotherm could bedetected in the primordial shoots with a water content belowabout 20%. When excised whole buds were cooled very slowly,the exotherm temperature shifted markedly to a lower value andthe exotherm became much smaller. Also, masses of needle icewere observed, mainly beneath the crown of the primordial shoot.From these results, it may be concluded that most of the waterin primordial shoots gradually migrates out through the crownand freezes as the temperature decreases (extraorgan freezing),which enables primordial shoots to survive at very low temperatures.Winter buds of Abies balsamea held at — 20?C for 30 daysand then slowly cooled down to —50 or —60?C remainedalive. Thus, there seems to be no low temperature limit to thisfrost avoidance mechanism, if the primordial shoots can resistintensive freeze-dehydration. Low temperature exotherms wereobserved in all genera which belong to Abietoideae and Laricoideaeof Pinaceae, all of which have a crown in the primordial shoots,but not in other conifers. ¹ Contribution No. 2037 from the Institute of Low TemperatureScience. (Received June 25, 1979; )     

Stand characteristics in colour-infrared aerial photographs as indicators of epiphytic lichens

Information about forest biodiversity has so far been collected mostly by using field inventories, but it is desirable to find other methods that can cover large areas at a lower cost. In a forest landscape covering 2000 ha in southern Sweden we tested if colour-infrared (CIR) aerial photographs on the scale of 1:30000 can be used to interpret forest stand characteristics correlated to the occurrence of epiphytic lichens that are Red-listed or otherwise indicate high nature conservation value (signal species). Using logistic regression we found that the interpreted stand variables tree height and crown structure class were significantly correlated to the occurrence of Red-listed species. For signal species, the variables tree height, percentage of southern deciduous trees and crown structure class were significantly correlated to the occurrence. The logistic regression models successfully predicted a significantly higher probability to find Red-listed species in the stands that actually contained such species compared to stands without Red-listed species. The same was true for stands with signal species. We conclude that interpretation of CIR aerial photographs could be a useful method to find certain groups of epiphytic lichens in surveys covering large areas.     

Effects of epiphytic lichens on host preference of the vascular epiphyte Tillandsia usneoides

We investigated the potential for nonvascular epiphytic species (primarily lichens) to affect the quality of different host tree species for the vascular epiphyte Tillandsia usneoides in the southeastern USA. Different host tree species had substantially different abundances of Tillandsia , and these abundances were correlated with the composition of nonvascular epiphyte communities. In greenhouse experiments Tillandsia grew significantly faster on the branches of Quercus virginiana (a species with very high natural abundances of Tillandsia ) when the dominant lichen on Q. virginiana was left intact than when the lichen was removed from the branches. In laboratory experiments, extracts from Cryptothecia rubrocincta , a lichen that was 10 times more common on poor host species for Tillandsia than on good host species, reduced Tillandsia seedling survival and growth in comparison to extracts from other species and rainwater. In field experiments, lichens increased the proportion of Tillandsia seeds and vegetative strands that adhered to the trunk of Ilex opaca (a poor Tillandsia host), but lichens did not affect propagule adherence to Q. virginiana . Our results are by no means exhaustive of the possibilities, but they suggest that the structure and diversity of vascular and nonvascular epiphytic communities that grow in different tree species may not be simply the product of host tree characteristics, but may also be influenced by interactions among the epiphytes themselves.     

Water relations and carbon dioxide exchange of epiphytic lichens in the Namib fog desert

Although there is only negligible rainfall, frequent nocturnal fog, dew and high air humidity support a luxurious lichen vegetation in the coastal zone of the central Namib Desert (Namibia). In earlier publications, we have studied ecophysiological performance of a series of epilithic and terrestrial lichens. Here, we have extended this work to three epiphytic species (Heterodermia namaquana, Ramalina lacera, and Xanthoria turbinata) that inhabit the sparse perennial shrubs growing in this area. Our intention, monitoring lichen CO₂ exchange, their water relations and microclimate conditions, was to determine the functional mechanisms that allow these epiphytes to exist under the special conditions of a fog desert. Measurements were conducted mainly during the spring season.The epiphytic lichens showed response patterns very similar to the epilithic and epigaeic species at the same site. Their metabolism was activated through moistening by dew and/or fog during the night and, in the very early morning, they exhibited the typical brief peak of net photosynthesis (NP) between sunrise and desiccation. The thalli were almost completely dry for the remainder of the day. Average duration of the positive NP during the morning peak was about 3 h. Dew condensation, alone, resulted in activation that provided 58–63% of integrated carbon income (ΣNP) as compared to fog (plus dew). In the late afternoon, there was a tendency for hydration to increase again, due to water vapour uptake at higher air humidity, and this allowed on some days a brief additional period of very low rates of photosynthesis shortly before sunset.Light response of photosynthesis showed “sun-plant” characteristics with saturation around 1000 μmol m⁻² s⁻¹ photosynthetically active photon flux density (PPFD). Light compensation point (LCP) of CO₂ exchange after sunrise was highly dependent on actual water content (WC) for X. turbinata: at low hydration it was ca. 10 μmol m⁻² s⁻¹ PPFD whilst, at high WC, it was almost 80 μmol m⁻² s⁻¹ PPFD. In contrast, LCP of R. lacera was almost independent of WC. This phenomenon was probably due to differences in thallus structure.Maximal attained NP and daily ΣNP both showed a saturation-type response to previous maximal nocturnal WC. Neither parameter was increased substantially when higher maximal thallus WCs were produced by experimental moistening in the night. All three species, despite their different morphologies, performed optimally at the highest nocturnal moistening achieved by natural fog and were not able to make use of higher hydration.The three studied epiphytes were similar in their chlorophyll-related rates of NP. Due to lower chlorophyll content, dry weight and carbon-related NP of X. turbinata was only about one-third of that of the other two species. The average carbon income on days with fog and/or dew hydration during the spring season amounted to 2.4 and 2.1 mgC (gC)⁻¹ day⁻¹ (related to thallus carbon content) for H. namaquana and R. lacera, respectively. This primary production was of similar magnitude to those found for the terrestrial species at the same site.     

Iron and phosphate uptake in epiphytic and saxicolous lichens differing in their pH requirements

The hypothesis is tested that pH-dependent Fe and P uptake influence the preference of epiphytic and saxicolous lichens for certain ranges of ambient pH. Five species from acidic substrata (Hypogymnia physodes, Parmeliopsis ambigua, and Platismatia glauca) or covering the range from weakly acidic to alkaline substrata (Lecanora muralis and Phaeophyscia orbicularis) were exposed to solutions of FeCl₂, FeCl₃, or KH₂PO₄ at pH 3 and 8 in the laboratory. Avoidance of alkaline substrata is explainable by low Fe³⁺ uptake at pH 8 in the case of H. physodes and the inability for net P uptake and membrane damage in P. ambigua at this pH. Preference for acidic substrata in Pl. glauca, however, is neither related to Fe nor P uptake. Efficient Fe³⁺ and P uptake at pH 8 explains the tolerance of L. muralis and Ph. orbicularis to alkaline conditions. Intracellular accumulation of Fe²⁺ in probably toxic amounts at pH 3 in Ph. orbicularis is correlated with the absence of this lichen from strongly acidic substrata. Avoidance of acidic sites by L. muralis is not attributable to Fe or P uptake. In summary, the results suggest that pH-dependent Fe and P uptake characteristics are involved in the determination of pH preferences of epiphytic and saxicolous lichens, but are not the only relevant factor.     

High cyanobiont selectivity of epiphytic lichens in old growth boreal forest of Finland

Here, cyanobiont selectivity of epiphytic lichen species was examined in an old growth forest area in Finland. Samples of the eight lichen species were collected from the same aspen (Populus tremula) and adjacent aspens in the same stand. The cyanobionts of these samples were compared with free and symbiotic Nostoc obtained from other habitats and geographic regions. Our results, based on the phylogenetic analysis of a partial small subunit of the ribosomal DNA (16S rDNA) and the rbcLX gene complex did not show any correlation with the geographic origin of the samples at any spatial scale. Instead, there was a correlation between the cyanobionts and the alleged taxonomy of their mycobionts. The results indicate that the lichen species examined are highly selective towards their cyanobiont partners. Only Lobaria pulmonaria proved to be more flexible, being able to associate with a wide range of Nostoc. A same Nostoc strain was found to form associations with taxonomically unrelated lichens indicating that the cyanobiont-mycobiont associations as a whole were not highly specific in the examined species.     

Distribution pattern of air pollution and epiphytic lichens in the Osaka Plain (Japan)

The geographic distributions of two common species of epiphytic lichens,Phaeophyscia limbata (Poelt) Kashiw. andLecanora pulverulenta Müll. Arg., on city park trees in the Osaka Plain and their relation to the distribution of SO₂ and NO₂ was examined. The concentrations of SO₂ and NO₂ are greater in the central area of this plain, and decrease with distance from the center. The frequency ofL. pulverulenta is lowest in the central area, and increases gradually toward the periphery. In contrast, the frequency ofP. limbata is highest in the intermediate area, and lower in the central and peripheral areas, i.e. at intermediate SO₂ and NO₂ concentrations. We conclude thatL. pulverulenta is more useful thanP. limbata as a bioindicator of air pollutants such as SO₂ and NO₂.     

Development of secondary woodland in oak wood pastures reduces the richness of rare epiphytic lichens

Wooded pastures with ancient trees were formerly abundant throughout Europe, but during the last century, grazing has largely been abandoned often resulting in dense forests. Ancient trees constitute habitat for many declining and threatened species, but the effects of secondary woodland on the biodiversity associated with these trees are largely unknown. We tested for difference in species richness, occurrence, and abundance of a set of nationally and regionally red-listed epiphytic lichens between ancient oaks located in secondary woodland and ancient oaks located in open conditions. We refined the test of the effect of secondary woodland by also including other explanatory variables. Species occurrence and abundance were modelled jointly using overdispersed zero-inflated Poisson models. The richness of the red-listed lichens on ancient oaks in secondary woodland was half of that compared with oaks growing in open conditions. The species-level analyses revealed that this was mainly the result of lower occupancy of two of the study species. The tree-level abundance of one species was also lower in secondary woodland. Potential explanations for this pattern are that the study lichens are adapted to desiccating conditions enhancing their population persistence by low competition or that open, windy conditions enhance their colonisation rate. This means that the development of secondary woodland is a threat to red-listed epiphytic lichens. We therefore suggest that woody vegetation is cleared and grazing resumed in abandoned oak pastures. Importantly, this will also benefit the vitality of the oaks.     

Changes in the distributions of epiphytic lichens in southern Sweden using a new statistical method

Past studies on changes in species distribution have mainly been based on analysis of range boundaries. In contrast, the method used here evaluates shifts in species' geographic centroids within a predefined area. We used presence/absence data on epiphytic lichens collected 1986 and 2003 from 64 sites in southern Sweden. A centroid was calculated each year, for each lichen species and substrate. The distance of centroid movement was evaluated in a permutation procedure. In total, 56 lichen species on 22 tree species were involved in the analyses, yielding 30 cases that had sufficient sample sizes both years to be evaluated. Out of these, three exhibited a significant movement of their centroid. The shift of lichen centroids of Hypogymnia physodes (L.) Nyl. and Vulpicida pinastri (Scop.) J.-E. Mattsson & M. J. Lai on the tree species Juniperus communis L. was 50 and 151 km with the direction 27° and 48°, respectively. For Hypogymnia physodes on Pinus sylvestris L., corresponding values were 41 km and 30°. The northnortheast shifts of these species in Sweden could be a response to a warming climate.     

Tree and stand-scale factors affecting richness and composition of epiphytic bryophytes and lichens in deciduous woodland key habitats

Conservation and sustainable forestry are essential in a multi-functional landscape. In this respect, ecological studies on epiphytes are needed to determine abiotic and biotic factors associated with high diversity. The aim of the present study was to evaluate relative sensitivity of conservation targets (epiphytic bryophytes and lichens) in relation to contrasting environmental variables (tree species, tree diameter at breast height, bark crevice depth, pH, tree inclination, pH, forest stand age, area and type) in boreo-nemoral forests. The study was conducted in Latvian 34 woodland key habitat (WKH) boreo-nemoral forest stands. Generalized linear mixed models and canonical correspondence analysis showed that tree species and tree bark pH were the most important variables explaining epiphytic bryophyte and lichen composition and richness (total, Red-listed, WKH indicator species). Forest stand level factors, such as stand size and habitat type, had only minor influence on epiphytic species composition and richness. The results of the present study indicate a need to maintain the diversity of tree species and large trees, particularly Acer platanoides, Carpinus betulus, Fraxinus excelsior, Populus tremula, Tilia cordata, Ulmus glabra and Ulmus laevis in conservation of epiphytic bryophyte and lichen communities in the future.     

Features of the biomass distribution of epiphytic lichens on scotch pine (Lower Angara Region)

The biomass of epiphytic lichens growing on Scots pine varies from 130 to 1090 g and is composed mainly of lichens from three genera: Bryoria (45%), Hypogymnia (34%), and Evernia (12%). The majority of lichens (66%) grow on tree branches (96%) in the zone of maximum development, located at a height of 9–13.5 m. The lichen biomass on tree trunks is insignificant (4%) and located mainly at their bottom part (70%).     

Evidence of high annual growth rate for lichens in the maritime Antarctic

On Livingston Island (South Shetland Islands, Antarctica) young moraines, now roughly 45 years old, were investigated in 1991 and 2002. More than 500 thalli of the 6 most abundant species, Acarospora macrocyclos, Bellemerea sp. Buellia latemarginata, Caloplaca sublobulata, Rhizocarpon geographicum and Usnea antarctica, previously measured in 1991, were measured again in 2002, which allowed an accurate measure of thallus growth rates. From our results, we believe it important to emphasize that: (1) lichen colonization probably took place at the end of the 1950s, soon after the last glacier retreat. (2) No relation between thallus growth rate and boulder size could be established. (3) Higher annual growth rates than those previously estimated were observed for R. geographicum and Bellemerea sp. and the former had one of the highest growth rates ever reported for this species. We speculate whether this high growth rate may be related to the rapid retreat of the Livingston Island ice cap and glaciers during the last recent years, and which have been attributed to climate warming on the Antarctic Peninsula region.     

Hierarchical analysis of the relationship between spatial distribution and abundance of epiphytic lichens (Mt. Olympos-Greece)

The relationship between distribution and abundance of epiphytic lichens on five different substrates (Fagus sylvatica, Pinus nigra, Quercus coccifera, Pinus heldreichii and Quercus pubescens) has been studied. For the first three substrates the above relationship has been studied hierarchically. It was concluded that this relationship is partly represented by the core-satellite distribution. This abundance-distribution pattern could probably be explained by more than one hypothesis and it is probably related with the life strategies of lichen species.     

Habitat structure and fish predation: effects on invertebrate colonisation and breakdown of stream leaf packs

1. We investigated the effects of two features of leaf‐pack habitat structure (i.e. mass of a leaf pack and surface area of leaves comprising a leaf pack) and fish predation on colonisation of shredders and leaf breakdown rates in a coldwater stream. Packs were constructed of red maple (Acer rubrum) leaves. 2. A 2 × 3 × 3 factorial experiment was used to manipulate fish predation (exclusion and control cage), leaf‐pack mass (1, 3 and 5 g dry mass) and leaf surface area (small: approx. 17.9 cm², medium: approx. 34.6 cm², large: approx. 65.6 cm²). Exclusion cages had mesh on all sides, whereas control cages lacked mesh on two sides to provide access to fish. 3. Common shredders were Gammarus pseudolimnaeus, Pycnopsyche and Lepidostoma. Shredder biomass per leaf pack increased with the mass of a leaf pack (P < 0.001), but biomass per unit mass of leaf pack did not differ with leaf‐pack mass (P = 0.506). Shredder densities did not respond to the exclusion of fish (P > 0.7) or leaf surface area (P > 0.7), and interactions among treatment factors were not significant (P > 0.2). 4. Breakdown rates were lower for leaf packs comprised of small leaves (P < 0.001) and leaf packs with high mass (P = 0.001). Excluding fish did not significantly affect leaf breakdown rates (P = 0.293), and interactions among treatment factors were not significant (P > 0.3). Breakdown rates were highest when packs consisted of few leaves (i.e. leaf packs with large leaves and low mass) and were colonised by many shredders. 5. Fish predation was not an important factor controlling shredder densities in leaf packs over the spatiotemporal scale of our experiment. Nevertheless, we found shredder colonisation was proportional to leaf‐pack mass and breakdown rates were affected by leaf‐pack size (i.e. number of leaves in a pack). We suspect that fragmentation is the primary mechanism causing the breakdown rates to be dependent on leaf‐pack size.     

Apple dwarfing rootstocks and interstocks affect the type of growth units produced during the annual growth cycle: precocious transition to flowering affects the composition and vigour of annual shoots

Background and Aims: Precocious flowering in apple trees is often associated witha smaller tree size. The hypothesis was tested that floral evocationin axillary buds, induced by dwarfing rootstocks, reduces thevigour of annual shoots developing from these buds comparedwith shoots developing from vegetative buds. Methods: The experimental system provided a wide range of possible treevigour using ‘Royal Gala’ scions and M.9 (dwarfing)and MM.106 (non-dwarfing) as rootstocks and interstocks. Second-yearannual shoots were divided into growth units corresponding toperiods (flushes) of growth namely, vegetative spur, extensiongrowth unit, uninterrupted growth unit, floral growth unit (bourse)and extended bourse. The differences between the floral andvegetative shoots were quantified by the constituent growthunits produced. Key Results: The dwarfing influence was expressed, firstly, in reduced proportionsof shoots that contained at least one extension growth unitand secondly, in reduced proportions of bicyclic shoots (containingtwo extension growth units) and shoots with an uninterruptedgrowth unit. In treatments where floral shoots were present,they were markedly less vigorous than vegetative shoots withrespect to both measures. In treatments with M.9 rootstock,vegetative and floral shoots produced on average 0·52and 0·17 extension growth units, compared with 0·77extension growth units per shoot in the MM.106 rootstock treatment.Remarkably, the number of nodes per extension growth unit wasnot affected by the rootstock/interstock treatments. Conclusions: These results showed that rootstocks/interstocks affect thetype of growth units produced during the annual growth cycle,reducing the number of extension growth units, thus affectingthe composition and vigour of annual shoots. This effect isparticularly amplified by the transition to flowering inducedby dwarfing rootstocks. The division of annual shoot into growthunits will also be useful for measuring and modelling effectsof age on apple tree architecture.