Different biomass-allocation patterns among four tree species in heavily disturbed sites on a volcanic mountain in Hokkaido,northern Japan

We have compared biomass-allocation patterns and frequency of sprouting among saplings of four tree species (Larix kaempferi, Betula platyphylla var. japonica, Populus maximowiczii, and Populus sieboldii) growing on a volcanic mountain in Hokkaido, northern Japan. Growing conditions were very harsh on the mountainside. Leaf mass and fine root mass relative to root mass were larger in L. kaempferi, and L. kaempferi root mass was less than for the other species. Sprouting ratios were high for the broadleaved species. Different allometries and sprouting ratios among species suggest that survival strategies for L. kaempferi were different from those for the broadleaved species. L. kaempferi has greater ability to increase leaf mass under harsh growing conditions; this probably results in large photosynthetic production by L. kaempferi on the volcano. In contrast, the two Populus species and Betula platyphylla seem to maintain populations through their ability to produce sprouts from large root systems.     

Chemical soil properties under five age series ofAlnus nepalensis plantations in the Eastern Himalayas

Summary Five age series stands ofAlnus nepalensis D. Don monocultures have been selected from the Pankhasari range of the Kalimpong forest division in the Eastern Himalayas. Chemical soil propeties such as organic carbon, total nitrogen, CN ratio, available phosphorus and exchangeable cations (Ca⁺², K⁺ and Na⁺) were analysed in all these different ages of plantation stands from the surface to 1 m deep profiles to find out the long term effect ofA. nepalensis on the soil quality of these erosion vulnerable slopes.Organic carbon, available phosphorus and total nitrogen content per-hectare increased with increasing plantation stand age. Analysis of variance for nutrients showed significant variation in depth (total nitrogenP<0.001; organic carbon,P<0.001; available phosphorus,P<0.05; exchangeable Ca⁺²,P<0.01; and exchangeable K⁺,P<0.01) and between plantation stands (organic carbon,P<0.05; and available phosphorusP<0.05). High value of total nitrogen content (34.97t ha^–1) was obtained in 46 yr old stand. Soil pH is low. It increased down the depth and in older plantations. These stands with high soil total nitrogen and organic carbon content show increased fertility of the stands. The cover ofA. nepalensis in rocky, landslide and ecosion prone slopes of the Eastern Himalayas, serves to protect and improve soil quality.     

Nitrate reductase and glutamine synthetase activities in relation to growth and nitrogen assimilation in red oak and red ash seedlings: effects of N-forms,N concentration and light intensity

The effects of growing seedlings of red oak (Quercus rubra) and red ash (Fraxinus pennsylvanica) with Hoagland solutions containing five N-regimes, differing in the N-forms (NH₄, NO₃) and concentrations (High and Low), in relation to light intensity were investigated by the utilization of enzymatic markers of the N assimilation pathway, nitrate reductase (NR) and glutamine synthetase (GS). Red oak and red ash showed different patterns of N-assimilation. Red oak seedlings assimilated NO₃ in low amounts in their roots and leaves, whereas red ash seedlings assimilated high amounts of NO₃, mostly in the leaves. A significant amount of constitutive NR activity was found in red oak seedlings supplied with NH₄ N-regime. This could be characteristic of a species adapted to soils that are poor in nitrogen. Root GS activity was lower in red oak seedlings than in red ash seedlings, indicating that the rate of NH₄ assimilation differed in these two hardwood species. Low irradiance reduced growth of both hardwood species, but greatly affected the specific leaf area of red ash and reduced NO₃ assimilation (when data are expressed per leaf area). Both species reacted similarly to N-regimes in terms of relative growth rate.     

Retardation of tree growth by injection of plant growth regulators

Previously topped American sycamore (Platanus occidentalis L.), cottonwood (Populus deltoides Bartr. ex Marsh), water oak (Quercus nigra L.), and bigleaf maple (Acer macrophyllum Pursh) trees were injected with aqueous solutions of maleic hydrazide or dikegulac-sodium in April-June, 1979 at various locations around the United States. Measurements made later in the autumn showed that dikegulac-sodium reduced sprout regrowth in all four species, whereas maleic hydrazide was only effective in controlling regrowth of American sycamore and cottonwood. The effects of dikegulac persisted through the second growing season. Significant variability was observed among treated trees and must be reduced in order to achieve consistent growth responses from an application of growth retardants.This paper reports the results of research only. Mention of a growth regulator does not consitute a recommendation for use by the U.S. Department of Agriculture nor does it imply registration under FIFRA as amended. Mention of a trademark or propriety product does not constitute a guarantee or warranty of the product by the U.S. Department of Agriculture and does not imply its approval to the exclusion of other products that may also be suitable.     

Effects of aluminium and mineral nutrition on growth and chemical composition of hydroponically grown seedlings of five different forest tree species

Forest die-back and impaired tree vitality have frequently been ascribed to Al-toxicity and Al-induced nutritional disorders due to increased acidification of forest soils. Therefore, in this experiment effects of Al were studied on growth and nutrient uptake with seedlings of five different forest tree species. During growth in culture solutions with and without Al all five species proved to be very Al-tolerant, despite high accumulation of Al in roots. In the coniferous evergreens Douglas-fir and Scots pine shoot as well as root Al concentrations were significantly higher than in the deciduous broad-leaved species oak and birch. Larch showed intermediate Al levels. In none of the five species did Al reduce nutrient concentrations or the Ca/Al ratio to values below the critical level. Besides differences in Al accumulation, coniferous and broad-leaved species also differed with respect to uptake and assimilation of nitrogen. Due to extra NH ₄ ⁺ uptake, oak and birch showed a much higher N uptake and higher NH ₄ ⁺ preference than the coniferous species. Especially with oak this high NH ₄ ⁺ preference in combination with a low specific root surface area resulted in a high root proton efflux density. In comparison to both broad-leaved trees and Scots pine the NO ₃ ⁻ reduction capacity of larch and Douglas-fir was extremely low. This may have important consequences for both species if grown in NO ₃ ⁻ -rich soils.     

Alnus rubra nodulation capacity of soil under five species from harvested forest sites in coastal British Columbia

Nodulation of Alnus rubra seedlings after inoculation with soil from under A. rubra, Betula papyrifera. Rubus lacianutus, R. spectabilis, and R.ursinus on 2 recently harvested sites was compared. Nodulation capacity was low compared to other published reports, ranging from 0 to 18.9 infective units cm^-3 of soil and was significantly affected by the site and plant species. Nodulation capacity of soil under alder was significantly higher than under all other species except R. spectabilis, regardless of site. The lowest nodulation capacity was found in soil under B. papyrifera.Joint appointment with Dept. of Soil Science, Faculty of Agricultural Sciences     

Effect of alginate matrix composition on regrowth of in vitro-derived encapsulated apical microcuttings of hybrid aspen

Effect of alginate matrix composition on regrowth performance of encapsulated microcuttings of hybrid aspen (Populus tremula L. × P. tremuloides Mincx.) was studied. Both high regrowth frequency and viability of explants were registered in all encapsulation mixtures tested. Some ingredients of the matrix (nutrient medium salts, sugars, growth regulators) significantly affected the initial development of the microcuttings. Sucrose appeared to play an important role in the starting stage of the regrowth event.     

3个温带阔叶树种根系长期分解速率研究

根系分解对土壤碳固持和养分长期有效性具有重要意义,但目前对根系的长期分解模式仍知之甚少。比较3个温带阔叶树种不同直径根系7 a分解动态,可为生态过程模型提供数据支撑。在帽儿山生态站采用分解袋法对白桦（Betula platyphylla）、春榆（Ulmus davidiana var.japonica）、水曲柳（Fraxinus mandshurica）的5个直径等级（< 1 mm）、（1-2 mm）、（2-5 mm）、（5-10 mm）、（10-20 mm）根系进行了7 a野外分解实验。重复测量方差分析表明：时间、树种、直径与树种交互作用、树种与时间交互作用显著影响根系质量保持率。根系质量保持率随时间呈指数下降趋势,7 a间根系经历早期快速分解和后期慢速分解两个阶段,实验结束时根系仍剩余相当部分的初始质量（24%-56%）。利用Olson指数衰减模型估计各树种根系分解系数发现：白桦根系分解系数与根直径间具有显著的线性正相关关系,水曲柳具有显著对数正相关关系,春榆的关系不显著。3个树种短期分解系数均高估根系分解速率,而且不能完全代表长期分解系数的种内、种间差异。研究结果对理解根系长期分解速率随直径变化模式提供了数据支撑。     

Gas composition and respiration of water oak (Quercus nigra L.) and green ash (Fraxinus pennsylvanica Marsh.) roots after prolonged flooding

Summary We compared the effects of 9.5 months of a continuous flooding treatment with a drained control treatment on one year-old seedlings of green ash (Fraxinus pennsylvanica Marsh.) and water oak (Quercus nigra L.), two tree species common to the bottomland-hardwood forests of eastern North America. The internal root gas composition of the more flood tolerant species, green ash, maintained higher oxygen and lower carbon dioxide concentrations under the flooding treatment than water oak. This apparently resulted in differences in rhizosphere oxidation. The amounts of Fe and Mn and the Fe/Mn ratio of the root coating extracted from trees in reduced soil conditions were much higher for the green ash than the water oak. It is argued that this reflects differences in the ability of these two species to maintain rhizosphere oxidation under prolonged periods of flooding and to prevent the accumulation of reduced potentially phytotoxic compounds. Alcohol dehydrogenase activity increased in the green ash and decreased in the water oak in the flooded treatment. This indicated that the better adapted species was able to rely upon increased anaerobic respiration in order to compensate for the decreased root oxygen supply, but the water oak was unable to maintain previous levels of respiration, probably as the result of sulfide toxicity.     

Effects of elevated CO<Subscript>2</Subscript> and nitrogen on wood structure related to water transport in seedlings of two deciduous broad-leaved tree species

Wood structure might be altered through the physiological responses to atmospheric carbon dioxide concentration ([CO₂]) and nitrogen (N) deposition. We investigated growth, water relations and wood structure of 1-year-old seedlings of two deciduous broad-leaved tree species, Quercus mongolica (oak, a ring-porous species) and Alnus hirsuta (alder, a diffuse-porous species and N₂–fixer), grown under a factorial combination of two levels of [CO₂] (36 and 72 Pa) and nitrogen supply (N; low and high) for 141 days in phytotron chambers. In oak, there was no significant effect of [CO₂] on wood structure, although elevated [CO₂] tended to decrease stomatal conductance (g _s) and increased water use efficiency regardless of the N treatment. However, high N supply increased root biomass and induced wider earlywood and larger vessels in the secondary xylem in stems, leading to increased hydraulic conductance. In alder, there was significant interactive effect of [CO₂] and N on vessel density, and high N supply increased the mean vessel area. Our results suggest that wood structures related to water transport were not markedly altered, although elevated [CO₂] induced changes in physiological parameters such as g _s and biomass allocation, and that N fertilization had more pronounced effects on non-N₂-fixing oak than on N₂-fixing alder.     

Drought acclimation of two deciduous tree species of different layers in a temperate forest canopy

Water-use strategies of Populus tremula and Tilia cordata, and the role of abscisic acid in these strategies, were analysed. P. tremula dominated in the overstorey and T. cordata in the lower layer of the tree canopy of the temperate deciduous forest canopy. Shoot water potential (), bulk-leaf abscisic acid concentration ([ABA]_leaf), abscisic acid concentration in xylem sap ([ABA]_xyl), and rate of stomatal closure following the supply of exogenous ABA (v) decreased acropetally through the whole tree canopy, and foliar water content per area (w), concentration of the leaf osmoticum (c), maximum leaf-specific hydraulic conductance of shoot (L), stomatal conductance (g_s), and the threshold dose per leaf area of the exogenous ABA (d_a) required to reduce stomatal conductance increased acropetally through the tree canopy (from the base of the foliage of T. cordata to the top of the foliage of P. tremula) in non-stressed trees. The threshold dose per leaf dry mass of the exogenous ABA (d_w) required to reduce stomatal conductance, was similar through the tree canopy. After a drought period (3 weeks), the , w, L, g_s, d_a and d_w had decreased, and c and v had increased in both species. Yet, the effect of the drought period was more pronounced on L, g_s, d_a, d_w and v in T. cordata, and on , w and c in P. tremula. It was concluded that the water use of the species of the lower canopy layer—T. cordata, is more conservative than that of the species of the overstorey, P. tremula. [ABA]_leaf had not been significantly changed in these trees, and [ABA]_xyl had increased during the drought period only in P. tremula. The relations between [ABA]_leaf, [ABA]_xyl and the stomatal conductance, the osmotic adjustment and the shoot hydraulic conductance are also discussed.     

Impact of tree species on nutrient and light availability: evidence from a permanent plot study of old-field succession

This paper compares vegetation composition, light availability, carbon and nutrient pools and Ellenberg indicator values among four old-field successional permanent plots that have received an initial treatment (ploughing, herbicide or sterilisation) prior to being left undisturbed in 1969, a second set of six plots received additional treatments (continued ploughing or mulching until 1982). On all plots species rich pioneer forests developed. Vegetation still varies among plots with different initial treatments: Sterilised plots can be distinguished from the others by dominance of Betula pendula, ploughed plots by Fraxinus excelsior, whereas herbicide-treated plots are intermediate with proportions of both species. By affecting light availability at the ground, tree species in turn influences ground vegetation and soil properties. Light availability turned out to be the dominant factor determining the composition of the vegetation in old-field succession.     

Evidence for light-independent and steeply decreasing PSII efficiency along twig depth in four tree species

Recent reports have indicated a considerably inactivated PSII in twig cortices, in spite of the low light transmittance of overlying periderms. Corresponding information for more deeply located and less illuminated tissues like xylem rays and pith are lacking. In this investigation we aimed to characterize the efficiency of PSII and its light sensitivity along twig depth, in conjunction with the prevailing light quantity and quality. To that aim, optical methods (spectral reflectance and transmittance, chlorophyll fluorescence imaging, low temperature fluorescence spectra) and photoinhibitory treatments were applied in cut twig sections of four tree species, while corresponding leaves served as controls. Compared to leaves, twig tissues displayed lower chlorophyll (Chl) levels and dark-adapted PSII efficiency, with strong decreasing gradients towards the twig center. The low PSII efficiencies in the inner stem were not an artifact due to an actinic effect of measuring beam or to an enhanced contribution of PSI fluorescence. In fact, the PSII/PSI ratios in cortices were higher and those in the xylem rays similar to that of leaves. Inner twig tissues were quite resistant to photoinhibitory treatments, tolerating irradiation levels several-fold higher than those encountered in their microenvironment. Moreover, the extent of high light tolerance was similar in naturally exposed and shaded twig sides. The results indicate an increasing, inherent and light-independent inactivation of PSII along twig depth. The findings are discussed on the basis of a recently proposed model for photosynthetic electron flow in twigs, taking into account the specific atmospheric and light microenvironment as well as the possible metabolic needs of such bulky organs.     

Carbon assimilation, nitrogen, and photochemical efficiency of different Himalayan tree species along an altitudinal gradient

In the area of Jumla region in Western Nepal, measurements of saturated leaf net photosynthetic rate (P_sat), nitrogen content, leaf fluorescence, carbon isotopic composition, and water status were performed on woody coniferous (Pinus wallichiana, Picea smithiana, Abies spectabilis, Juniperus wallichiana, Taxus baccata), evergreen (Quercus semecarpifolia, Rhododendron campanulatum), and deciduous broadleaved species (Betula utilis, Populus ciliata, Sorbus cuspidata) spreading from 2 400 m up to the treeline at 4 200 m a.s.l. With the exception of J. wallichiana, P_sat values were lower in coniferous than broadleaved species. Q. semecarpifolia, that in this area grows above the coniferous belt between 3 000 and 4 000 m, showed the highest P_sat at saturating irradiance and the highest leaf N content. This N content was higher and P_sat lower than those of evergreen oak species of tempe forests at middle and low altitudes. For all species, P_sat and N content were linearly correlated, but instantaneous nitrogen use efficiency was lower than values measured in lowland and temperate plant communities. The values of carbon isotopic composition, estimated by ₁₃C, showed the same range reported for temperate tree species. The ranking of ₁₃C values for the different tree types was conifers < evergreen broadleaved13C were found along the altitudinal gradient. Quantum yield of photochemistry at saturating irradiance, measured by leaf fluorescence (F/F_m), was highest in J. wallichiana and lowest in T. baccata. Overall, photochemical efficiency was more strongly related to species than to altitude. Interestingly, changes of .F/F_m along the altitudinal gradient correlated well with the reported altitudinal distribution of the species.This revised version was published online in March 2005 with corrections to the page numbers.     

Fine root architecture, morphology, and biomass of different branch orders of two Chinese temperate tree species

We have limited understanding of architecture and morphology of fine root systems in large woody trees. This study investigated architecture, morphology, and biomass of different fine root branch orders of two temperate tree species from Northeastern China—Larix gmelinii Rupr and Fraxinus mandshurica Rupr —by sampling up to five fine root branch orders three times during the 2003 growing season from two soil depths (i.e., 0–10 and.10–20 cm). Branching ratio (R _b) differed with the level of branching: R _b values from the fifth to the second order of branching were approximately three in both species, but markedly higher for the first two orders of branching, reaching a value of 10.4 for L. gmelinii and 18.6 for F. mandshurica. Fine root diameter, length, SRL and root length density not only had systematic changes with root order, but also varied significantly with season and soil depth. Total biomass per order did not change systematically with branch order. Compared to the second, third and/or fourth order, the first order roots exhibited higher biomass throughout the growing season and soil depths, a pattern related to consistently higher R _b values for the first two orders of branching than the other levels of branching. Moreover, the differences in architecture and morphology across order, season, and soil depth between the two species were consistent with the morphological disparity between gymnosperms and angiosperms reported previously. The results of this study suggest that root architecture and morphology, especially those of the first order roots, should be important for understanding the complexity and multi-functionality of tree fine roots with respect to root nutrient and water uptake, and fine root dynamics in forest ecosystems.     

Spatial patterns of tree seedling establishment and their relationship to environmental variables in a cold-temperate deciduous forest of eastern North America

Patterns of seedling recruitment may have persistent effects on population and community processes. Assuming seed availability is not limiting, the environmental sieve (i.e., the suite of factors influencing seed germination and seedling emergence and survival) determines how many seedlings establish and, most importantly, where they do so. In this study, we identify the spatial structure of some resources and abiotic conditions known to be significant for tree seedling emergence and survival and determine how these environmental factors influence the establishment of Fagus grandifolia, Acer saccharum, Fraxinus americana, and Ostrya virginiana in a deciduous forest of southern Québec (Canada). We expect an increase from Fagus, through Acer and Fraxinus, to Ostrya in the control of environmental variables on seedling emergence and survival, because of differences in the seed size of these species. Density of newly-emerged seedlings of all four species showed positive spatial autocorrelation at distances of up to ca. 10 m. Environmental variables were also structured at the same spatial scale, except for soil moisture. Acer seedling emergence pattern was positively correlated to photosynthetic photon flux density (PPFD), and the pattern of Fraxinus to soil N and moisture. Seedling survival was not spatially autocorrelated for any of the four species, although it was positively density-dependent in Acer and Fagus. In only Ostrya was seedling survival correlated (positively) to one of the environmental variables studied, i.e., PPFD. Overall, environmental variables were spatially less heterogeneous than seedling emergence and survival. Either seed availability was not saturating or factors not considered here, such as competition and predation (the intensity of which often varies with resources and/or abiotic conditions), modified the influence that the physical environment had on patterns of seedling establishment. Our prediction of a greater environmental control on seedling emergence and survival in small-seed species was not totally confirmed.     

北京典型绿化树种幼苗光合特性对硬化地表的响应

以北京典型绿化树种油松(Pinus tabulaeformis)、白蜡(Fraxinus chinensis)和元宝枫(Acer truncatum)幼苗为试验材料,设置透水硬化地表、不透水硬化地表和对照3个水平的地表类型,通过监测地表温度、土壤含水量(20 cm)和光合参数等指标,分析3种树木叶片光合特性对硬化地表的响应特征。结果表明:城市硬化地表极显著地提高了地表温度(P0.01),表现为透水地表不透水地表对照;不透水地表极显著地降低了3种树木土壤含水量(P0.01);透水地表极显著地提高了油松和元宝枫土壤含水量(P0.01)。两种硬化地表下油松、元宝枫和不透水地表下白蜡幼苗叶片的净光合速率、蒸腾速率和气孔导度降低,株高和基径生长受到抑制。硬化地表对油松和元宝枫的影响主要源于硬化地表形成的高温胁迫,对白蜡的影响主要源于不透水地表形成的干旱胁迫。因此,在配置和种植城市绿化植物时,为缓解硬化地表对城市树木生长发育的不利影响,应选择适应高温和干旱的植物,并采取降温和灌溉措施来降低硬化地表的影响。     

固氮树种旱冬瓜对退化林地土壤修复和林下植被重建的生态驱动效应

固氮树种对退化土壤和森林的结构与功能修复和改良具有重要价值,但是,固氮树种的混交配置、林龄变化和人工与天然起源林分对土壤修复和林下植被重建方面的生态驱动效应与影响机制的认识尚不够深入。因此,在金沙江上游选择退耕地上不同混交配置模式、不同林龄旱冬瓜(Alnus nepalensis)人工林和旱冬瓜天然次生林,对比分析了土壤养分、林下植被生长、林下植被多样性和旱冬瓜材积等指标的差异,以期阐明固氮树种旱冬瓜对土壤养分状况及林下植被重建的生态驱动效应。结果表明：(1)不同旱冬瓜人工林配置模式中,旱冬瓜纯林的土壤养分综合指标显著高于旱冬瓜+云南松混交林和云南松纯林(P<0.05);(2)不同林龄旱冬瓜人工林中土壤有机质、总氮及土壤养分综合指标呈现8年生林均显著大于5年生林(P<0.05);(3)土壤有机质、总氮及土壤养分综合指标均为旱冬瓜天然林>旱冬瓜和云南松人工混交林>云南松天然林(P<0.05);(4)SEM模型分析发现,旱冬瓜材积与土壤养分综合指标呈极显著的正相关关系(R=0.473,P<0.001);(5)在不同旱冬瓜人工配置模式和不同优势种的天然林中...     

尾叶桉林下5种植物的耐阴性、生物量及其个体消长

为了解尾叶桉林下不同植物种类的生理生态特性和生长适应性,在鹤山站尾叶桉林中选取分布较多且药用价值和经济价值较高的5种林下植物为研究对象,测定各植物的叶片特征、叶绿素含量和光合-光响应曲线,同时结合2006年和2011年尾叶桉林的样方调查数据分析了5种林下植物生物量的变化和个体数量的消长。结果表明:(1)野牡丹叶片的宽度、厚度、单片叶面积等指标最大,梅叶冬青最小,桃金娘、栀子和山苍子处于两者之间;桃金娘、野牡丹和山苍子的光合速率随光照的增强迅速增大,其光饱和点较高;梅叶冬青和栀子的光合速率也随光照的增强而增大,但其斜率比野牡丹和山苍子大,并很快就达到光饱和点。综合聚类分析植物的耐阴程度,5种植物可分为2类:第1类为耐阴性较强的梅叶冬青、栀子,第2类为耐阴性较弱的野牡丹、山苍子和桃金娘。(2)随着尾叶桉林的生长,5种林下植物的生物量都有不同程度的增加,其中增幅较大的是野牡丹和桃金娘分别增加74倍和18倍,增幅最小的是栀子仅1.3倍。野牡丹、桃金娘和山苍子这3个种生物量的增加主要是由个体高度和地径的增长引起,而栀子和梅叶冬青生物量的增加是其个体数量、株高和地径做出的贡献。(3)在尾叶桉林生长过程中,耐阴种类梅叶冬青随着个体密度、株高和地径的增长,其在林内发展成为主要优势灌木种类;栀子的密度虽然有一定的增大,但其株高和地径增加量小,生物量也小,发展为次要灌木种类。野牡丹、桃金娘和山苍子等3种的个体密度都在下降,虽然其地径、株高和生物量也有一定的增长,但其较强的阳生性不能适应林内的荫蔽环境,只适于林窗或林缘光照较强的地方生长。因此,在桉树林经营管理过程中应适当保护和种植梅叶冬青和栀子;在短轮伐期经营模式下,林窗和林缘也可适当保留野牡丹、桃金娘和山苍子。     

Chemical variation in two species of theLecanora subfusca group (Lecanoraceae,lichenizedAscomycotina)

The chemical variation inLecanora epibryon andL. subimmersa, two species of theL. subfusca group, has been examined. In both species the chemical differences are correlated with geographical distribution, but not with morphological differences. As a consequence the chemotypes are recognized at subspecific level. InL. epibryon three chemical races are segregated according to the various chemosyndromes present. Subspeciesepibryon, containing atranorin and triterpenoids, occurs in the northern hemisphere and South America, while the other two subspecies occur only in the southern hemisphere. The subsp.broccha (Nyl.)Lumbsch, comb. nova, contains atranorin, the stictic acid and the 2,5,7-trichloro-3-O-methylnorlichexanthone chemosyndromes, while subsp.xanthophora Lumbsch, subsp. nova, is similar but lacks the stictic acid chemosyndrome. Two chemical races occur in the pantropical speciesL. subimmersa. While subsp.subimmersa contains atranorin and zeorin, subsp.ramboldii Lumbsch & Elix, subsp. nova, contains an additional ten chlorinated xanthones.L. impressa is reduced to synonymy toL. subimmersa.