Carbon-based Secondary Compounds Research Articles

Intra-specific variation in lichen secondary compounds across environmental gradients on Signy Island, maritime Antarctic

Lichens produce various carbon-based secondary compounds (CBSCs) in response to abiotic conditions and herbivory. Although lichen CBSCs have received considerable attention with regard to responses to UV-B exposure, very little is known about intra-specific variation across environmental gradients and their role in protection against herbivory in the Antarctic. Here we report on the variation in CBSCs of two widely distributed and common Antarctic lichens, Usnea antarctica and Umbilicaria antarctica, between sites with different solar exposure (NW–SE) and along natural nitrogen (N) gradients which are associated with changing lichen-invertebrate associations on Signy Island (South Orkney Islands, maritime Antarctic). Fumarprotocetraric and usnic acid concentrations in Usnea showed no relationships with solar exposure, lichen-N or associated invertebrate abundance. However, fumarprotocetraric acid concentration was 13 times higher at inland sites compared to coastal sites along the N-gradients. Gyrophoric acid concentration in Umbilicaria was 33% lower in sun-facing (northerly exposed) habitats compared to more shaded (south-facing) rocks and declined with elevation. Gyrophoric acid concentration was positively correlated with the abundance and species richness of associated microarthropods, similar to the patterns found with lichen N. This initial investigation indicates that there can be large intraspecific variation in lichen CBSC concentrations across relative short distances (< 500 m) on Signy Island and raises further questions regarding current understanding of the role of CBSCs in Antarctic lichens in relation to biotic and abiotic pressures.

Contrasting responses of plant and lichen carbon‐based secondary compounds across an elevational gradient

Abstract Vascular plants and lichens often produce a diversity of carbon‐based secondary compounds (CBSCs) to protect them against biotic and abiotic stresses. These compounds play important but often compound‐specific roles in community and ecosystem processes by affecting herbivore and decomposer activity. However, our understanding of what drives community‐level CBSCs among ecosystems or across environmental gradients is limited. We measured concentrations and compositions of CBSCs for all dominant vascular plant and lichen species present across a 500‐m alpine elevational gradient. These measurements were combined with data on species composition and abundance to obtain whole‐community measures of plant and lichen CBSCs across the gradient. At the whole community level, plant CBSCs had the lowest concentrations while lichen CBSCs had the highest concentrations at the highest elevations. Further, plant CBCSs shifted from those associated with herbivore defence towards those protecting against light and oxidative stress as elevation increased, while lichen CBSCs showed the opposite pattern. Synthesis. Our findings that individual compounds show contrasting responses to the same environmental gradient highlight the importance of studying qualitative as well as quantitative changes in CBSCs. Further, the divergent responses between vascular plants and lichens reveal that in systems where both groups are abundant, they need to be considered simultaneously to better understand how future environmental changes may impact on ecosystem‐level processes. A free Plain Language Summary can be found within the Supporting Information of this article.

Gastropod grazing of epiphytic lichen-dominated communities depends on tree species

Tree species differ in longevity, canopy structure, and bark texture, chemistry and water storage. Tree species-specific traits play a role in shaping epiphytic vegetation and likely influence the community assembly of organisms feeding on epiphytes. Lichenivorous gastropods, species with calcium-rich shells in particular, need calcium and likely occur more abundantly in and around tree species with high available calcium. We quantified gastropod grazing on the epiphytic lichens Lobaria pulmonaria and Lobaria scrobiculata transplanted to blocks of adjacent trunks of Acer platanoides, Quercus robur and Tilia cordata. We tested the hypothesis that tree species known to have more available Ca, exhibit more grazing damage on transplanted lichens than trees with lower Ca-availability. The grazing pressure was 1.6–1.8 times higher for lichen transplants on Acer and Tilia known to produce litter with easily soluble Ca than on Quercus, which binds Ca as oxalate. Trees with a high grazing pressure on transplants had greater natural abundance of Lobaria virens than of L. pulmonaria. Gastropods preferred L. scrobiculata to L. pulmonaria, evidenced by more observed grazing marks and greater measured biomass loss. We attribute this difference to the lower concentration of carbon-based secondary compounds in L. scrobiculata. However, the strength of the preference varied between the three tree species receiving lichen transplants and was strongest on A. platanoides, while gastropods on T. cordata grazed equal amounts of each transplanted lichen. In conclusion, tree species influenced grazing patterns of gastropods on epiphytic lichens. In addition to bark pH and other factors, we have shown that tree species-specific differences in grazing pressure play a role in shaping the epiphytic macrolichen community.

Successful Colonization of Lodgepole Pine Trees by Mountain Pine Beetle Increased Monoterpene Production and Exhausted Carbohydrate Reserves.

Lodgepole pine (Pinus contorta) forests have experienced severe mortality from mountain pine beetle (MPB) (Dendroctonus ponderosae Hopkins) in western North America for the last several years. Although the mechanisms by which beetles kill host trees are unclear, they are likely linked to pine defense monoterpenes that are synthesized from carbohydrate reserves. However, how carbohydrates and monoterpenes interact in response to MPB colonization is unknown. Understanding this relationship could help to elucidate how pines succumb to bark beetle attack. We compared concentrations of individual and total monoterpenes and carbohydrates in the phloem of healthy pine trees with those naturally colonized by MPB. Trees attacked by MPB had nearly 300% more monoterpenes and 40% less carbohydrates. Total monoterpene concentrations were most strongly associated with the concentration of sugars in the phloem. These results suggest that bark beetle colonization likely depletes carbohydrate reserves by increasing the production of carbon-rich monoterpenes, and other carbon-based secondary compounds. Bark beetle attacks also reduce water transport causing the disruption of carbon transport between tree foliage and roots, which restricts carbon assimilation. Reduction in carbohydrate reserves likely contributes to tree mortality.

Carbon-based secondary compounds in the lichen Hypogymnia physodes deter detrivorous woodlice

Abstract Woodlice are not widely recognized as lichen-feeding invertebrates. We sought to discover whether the woodlouse, Porcellio scaber, could feed on the lichen Hypogymnia physodes and if the lichen carbon-based secondary compounds CBSCs would reduce grazing. We cut lichen thalli in two pieces, one was non-destructively rinsed in acetone to remove CBSCs; the other served as a control. Both pieces were fed to woodlice in a choice experiment. The CBSC concentration of individual thalli ranged from 3 to 19%. The woodlice grazed all pieces, but preferred the acetone-rinsed pieces, depending on the amount of CBSCs present in the non-extracted counterpart. The woodlice were not deterred from feeding on samples with CBSC concentrations ≤5%, which corresponded to natural contents in shade-adapted thalli. This suggested that P. scaber tolerates this amount of compounds at least in the short-term. In conclusion, P. scaber can feed on H. physodes, but CBSCs deter them from feeding.

Genomewide transcriptional reprogramming in the seagrass Cymodocea nodosa under experimental ocean acidification.

Here, we report the first use of massive-scale RNA-sequencing to explore seagrass response to CO2 -driven ocean acidification (OA). Large-scale gene expression changes in the seagrass Cymodocea nodosa occurred at CO2 levels projected by the end of the century. C.nodosa transcriptome was obtained using Illumina RNA-Seq technology and de novo assembly, and differential gene expression was explored in plants exposed to short-term high CO2 /low pH conditions. At high pCO2 , there was a significant increased expression of transcripts associated with photosynthesis, including light reaction functions and CO2 fixation, and also to respiratory pathways, specifically for enzymes involved in glycolysis, in the tricarboxylic acid cycle and in the energy metabolism of the mitochondrial electron transport. The upregulation of respiratory metabolism is probably supported by the increased availability of photosynthates and increased energy demand for biosynthesis and stress-related processes under elevated CO2 and low pH. The upregulation of several chaperones resembling heat stress-induced changes in gene expression highlighted the positive role these proteins play in tolerance to intracellular acid stress in seagrasses. OA further modifies C.nodosa secondary metabolism inducing the transcription of enzymes related to biosynthesis of carbon-based secondary compounds, in particular the synthesis of polyphenols and isoprenoid compounds that have a variety of biological functions including plant defence. By demonstrating which physiological processes are most sensitive to OA, this research provides a major advance in the understanding of seagrass metabolism in the context of altered seawater chemistry from global climate change.

Simulated global warming increases usnic acid but reduces perlatolic acid in the mat-forming terricolous lichenCladonia stellaris

AbstractLichens as sessile and slow-growing symbiotic associations have evolved various carbon-based secondary compounds (CBSCs) to mitigate the effects of some stressors in the extreme environments in which they often grow. The mat-forming lichenCladonia stellaris, an important fodder for reindeer, produces usnic acid in the outermost layer and perlatolic acid in the medulla. Here, we studied effects of simulated global warming on these CBSCs inC. stellariscultivated in climate chambers with: 1) ambient conditions as control or 2) ambient conditions +4°C. The chambers simulated, at an hourly resolution, an averaged 10-year growing season dynamic from a long-term monitored boreal mire in northern Sweden. After two months of acclimation, +4°C warming in one simulated growing season increased the concentration of usnic acid by 31% compared with ambient conditions, whereas the warming decreased the concentration of perlatolic acid by 14%. Since lichen CBSCs play important roles in ecosystem processes such as lichenivory and decomposition, these changes may profoundly affect lichen-dominated ecosystems.

Fungal tissue per se is stronger as a UV-B screen than secondary fungal extrolites in Lobaria pulmonaria

Abstract To test the hypotheses that (1) protective mycobiont tissues and/or (2) medullary UV-B-absorbing carbon-based secondary compounds (CBSCs) protect lichen photobionts against UV-B radiation, we quantified cortical UV-transmittance and ran a three-way factorial lab experiment with (1) three UV radiation regimes, (2) photobiont layers with/without a screening cortex, and (3) with natural/reduced CBSC-concentration. We used melanin-deficient Lobaria pulmonaria from shaded forests. Maximum photochemical efficiency of photosystem II ( F v/ F m) in photobionts inside thalli with natural CBSC-concentrations was not affected by any UV-regime, consistent with close to 0% measured cortical transmittance of wavelengths P P P = 0.001), mainly affecting exposed photobiont layers given short-wavelength UV radiation. Based on these findings, we conclude that the primary role of extrolites in L. pulmonaria is not to screen excess solar radiation.

Increased snow accumulation reduces survival and growth in dominant mat-forming arctic-alpine lichens

AbstractRelative growth rates (RGR) and carbon-based secondary compounds (CBSCs) were quantified in four dominant terricolous arctic-alpine mat-forming lichens with different preferences for snow cover. The aim was to evaluate the effects of snow depth, and thus snow cover duration, on lichen growth and performance. The species,Alectoria ochroleuca,Flavocetraria nivalis,Cladonia mitisandCetrariella delisei, are associated with increasing snow depth, respectively. They were transplanted for one year at five snow depths (0, 60, 120, 160 and 200 cm measured in early May) along each of four natural ridge to snow bed gradient transects in oceanic-alpine sites (western Norway). Snow slightly thicker than in source habitats caused negative RGR in the ridge top-dependentA. ochroleucaand the co-occurringF. nivaliswith somewhat higher snow tolerance. OnlyC. mitiswith the broadest ecological niche had positive RGR along most of the gradients (0–160 cm), even outside its natural range. The most snow-tolerant species,C. delisei, tolerant also to temporal inundation in water, had the lowest RGR. Nevertheless, it performed as well in places with little or no snow as in places where it grows naturally. Snow depth significantly affected total concentrations of CBSCs mainly inA. ochroleuca, which experienced substantial mass loss under snow. There was a highly significant increase in usnic acid concentration with increasing mass loss inA. ochroleuca, probably because usnic acid decays more slowly than other components. In conclusion, snow reduced lichen RGR, but in species-specific ways. Therefore, increasing snow depthper sealong spatial and/or temporal scales likely reduces abundance and distribution of dominant mat-forming lichens in the alpine ecosystems of Scandinavia.

The role of fungal parasites in tri-trophic interactions involving lichens and lichen-feeding snails.

Lichens are hosts for a variety of lichenicolous fungi. By investigating two lichens with specialized parasites, we will test the hypothesis that these parasites reduce lichen fitness by increasing the palatability of their respective hosts. The palatability of Lobarina scrobiculata and Lobariapulmonaria with or without galls of the lichenicolous fungi, Plectocarpon scrobiculatae and P.lichenum, respectively, were quantified in a feeding-preference experiment with grazing snails (Cepaea hortensis). We repeated the experiment for pairs with or without gall in which the carbon-based secondary compounds (CBSCs) had been reduced nondestructively by acetone rinsing. Lichens with galls had lower concentration of CBSCs than those without, but this contrast disappeared after acetone rinsing. In the lichen high in nitrogen (N) (the cyanolichen L.scrobiculata), the grazing was low, and the snails did not discriminate between specimens with and without Plectocarpon-galls. In L.pulmonaria low in N (green algae as main photobiont), the parasite reduced the lichen C:N ratio and the snails strongly preferred specimens with Plectocarpon-galls, regardless of whether CBSC concentration had been reduced or not. In conclusion, some lichen parasites can indirectly reduce lichen fitness by increasing its palatability and thus the grazing pressure from snails, whereas other parasites do not affect grazing preferences.

Effect of light intensity and wavelength on concentration of plant secondary metabolites in the leaves of Flourensia cernua

Flourensia cernua (tarbush) is a shrub that has encroached into grasslands in many areas of the northern Chihuahuan Desert and contains high levels of carbon-based secondary compounds. Concentrations of secondary compounds are affected by numerous biotic and abiotic influences, including amount and wavelength of solar radiation. However, responses to shade and ultraviolet light restriction are inconsistent among plant species and compound class. We conducted a three-year study to evaluate the effect of shade and UV light restriction on total phenolic and terpene concentrations in tarbush. Sixty plants were randomly assigned to one of three treatments (control, UV light restriction, or 50% incident light restriction). Mean concentrations of total phenolics and total volatiles in tarbush were 82.4 and 12.5 mg/g DM, respectively. Total phenolics did not differ between UV-restricted and control plants, but were lower in shaded plants than the other treatments (P < 0.05). Total volatiles tended to be greater for the UV-restricted treatment than control plants (P = 0.056), with shaded plants not different from either treatment. Treatment effects were detected for 18 individual compounds (P < 0.05). Our results partially support the hypothesis that UV restriction and shading alter carbon-based secondary chemical concentrations.

Carbon based secondary compounds do not provide protection against heavy metal road pollutants in epiphytic macrolichens

Lichens are useful monitoring organisms for heavy metal pollution. They are high in carbon based secondary compounds (CBSCs) among which some may chelate heavy metals and thus increase metal accumulation. This study quantifies CBSCs in four epiphytic lichens transplanted for 6months on stands along transects from a highway in southern Norway to search for relationships between concentrations of heavy metals and CBSCs along a gradient in heavy metal pollutants. Viability parameters and concentrations of 21 elements including nutrients and heavy metals in these lichen samples were reported in a separate paper. Medullary CBSCs in fruticose lichens (Ramalina farinacea, Usnea dasypoga) were reduced in the most polluted sites, but not in foliose ones (Parmelia sulcata, Lobaria pulmonaria), whereas cortical CBSC did not change with distance from the road in any species. Strong positive correlations only occurred between the major medullary compound stictic acid present in L. pulmonaria and most heavy metals, consistent with a chelating role of stictic acid, but not of other studied CBSCs or in other species. However, heavy metal chelating did not protect L. pulmonaria against damage because this species experienced the strongest reduction in viability in the polluted sites. CBSCs with an accumulation potential for heavy metals should be quantified in lichen biomonitoring studies of heavy metals because they, like stictic acid, could overshadow pollutant inputs in some species rendering biomonitoring data less useful. In the two fruticose lichen species, CBSCs decreased with increasing heavy metal concentration, probably because heavy metal exposure impaired secondary metabolism. Thus, we found no support for a heavy metal protection role of any CBSCs in studied epiphytic lichens. No intraspecific relationships occurred between CBSCs versus N or C/N-ratio. Interspecifically, medullary CBSCs decreased and cortical CBSCs increased with increasing C/N-ratio.

Removal of secondary compounds increases invertebrate abundance in lichens

Abstract We investigated how lichen carbon-based secondary compounds (CBSCs) affect abundance of invertebrates in five lichen species growing on the forest floor ( Cladonia rangiferina , Cladonia stellaris ) or on tree trunks ( Evernia prunastri, Hypogymnia physodes, Pseudevernia furfuracea ). To do this, CBSCs were removed by rinsing lichen thalli in acetone (which has no adverse effects on the lichens) and the lichens were re-transplanted in their natural habitat. After 4 months there was higher abundance of mites, springtails and spiders in the three epiphytic lichens that had their CBSC concentrations reduced. The increase in predatory spiders following CBSC reduction suggests that the compounds have multitrophic consequences. The acetone treatment reduced the number of nematodes in four of the lichen species. Given that lichens serve as important habitats for a diverse range of invertebrates, increased knowledge of how lichen CBSCs may regulate their abundance helps us to better understand the role that lichens and their defence compounds play in structuring forest food webs.

Lichen physiological traits and growth forms affect communities of associated invertebrates.

While there has been much interest in the relationships between traits of primary producers and composition of associated invertebrate consumer communities, our knowledge is largely based on studies from vascular plants, while other types of functionally important producers, such as lichens, have rarely been considered. To address how physiological traits of lichens drive community composition of invertebrates, we collected thalli from 27 lichen species from southern Norway and quantified the communities of associated springtails, mites, and nematodes. For each lichen species, we measured key physiological thallus traits and determined whether invertebrate communities were correlated with these traits. We also explored whether invertebrate communities differed among lichen groups, categorized according to nitrogen-fixing ability, growth form, and substratum. Lichen traits explained up to 39% of the variation in abundances of major invertebrate groups. For many invertebrate groups, abundance was positively correlated with lichen N and P concentrations, N:P ratio, and the percentage of water content on saturation (WC), but had few relationships with concentrations of carbon-based secondary compounds. Diversity and taxonomic richness of invertebrate groups were sometimes also correlated with lichen N and N:P ratios. Nitrogen-fixing lichens showed higher abundance and diversity of some invertebrate groups than did non-N-fixing lichens. However, this emerged in part because most N-fixing lichens have a foliose growth form that benefits invertebrates, through, improving the microclimate, independently of N concentration. Furthermore, invertebrate communities associated with terricolous lichens were determined more by their close proximity to the soil invertebrate pool than by lichen traits. Overall, our results reveal that differences between lichen species have a large impact on the invertebrate communities that live among the thalli. Different invertebrate groups show contrasting responses to traits that are indicative of thallus quality (nutrient concentrations), and thallus growth form is often an important determinant of the invertebrate community. Given the large diversity of lichen traits and growth forms that occur in many ecosystems, lichen-invertebrate communities may be an important contributor to overall community diversity in boreal forests.

Relative growth rates and secondary compounds in epiphytic lichens along canopy height gradients in forest gaps and meadows in inland British Columbia

We explore relative growth rates (RGRs) and carbon-based secondary compounds (CBSCs) in epiphytic lichens along height-above-the-ground gradients. The chlorolichen (Letharia vulpina (L.) Hue), the cephalolichen (Lobaria pulmonaria (L.) Hoffm.), and the cyanolichens (Lobaria hallii (Tuck.) Zahlbr., Nephroma helveticum Ach.) were attached to branches at 0.5–3.0 m heights of young spruce trees transplanted for 1 year in forest gaps and in old meadows of an inland valley in British Columbia. Cephalolichen and cyanolichen RGRs were highest in forest gaps, whereas the chlorolichen grew faster in meadows with twice as much light as forest gaps. Transplantation height did not influence lichen temperature or RGRs, despite height-dependent light increases. CBSCs were highest in the chlorolichen (13%), followed by the cephalolichen (5%) and the cyanolichens (1% and 0%). CBSC concentrations increased with thallus size, and were significantly higher in forest gaps mainly for L. pulmonaria. Only one minor CBSC in each species varied with height. The slow growth of cephalolichen and cyanolichens in meadows is consistent with these lichens’ preferences for forested habitats. Cold air ponding from snow-capped mountains was probably strong enough to form enough nocturnal dew to support reasonable lichen growth at all heights. The high species-specific and the low habitat-specific CBSC variations are consistent with constitutive CBSC defense levels in studied lichens.

Do lichen secondary compounds play a role in highly specific fungal parasitism?

Abstract Chemical interactions between highly host-specific lichenicolous fungi and their lichen hosts have been little studied. In an allometric study, we quantified carbon-based secondary compounds (CBSCs) in a mixed natural Lobarina scrobiculata population (N = 147) of the normal and the stictic acid-deficient chemotypes, both with and without galls of Plectocarpon scrobiculatae. We assessed the correlation between the presence/abundance of parasite galls and the lichen CBSCs contents, and quantified size-dependent contents of CBSCs. The parasite produced galls similarly in both chemotypes, indicating that the stictic acid complex does not deter Plectocarpon. Within both chemotypes, thalli with Plectocarpon had half the contents of all individual CBSCs than those without galls. There was a significant size-dependent increase in CBSC contents in thalli without galls, but not in those with. This study shows that lichen chemistry is involved in highly host-specific fungal parasitism, and widens our knowledge of specialized biotrophic fungal interactions.

Growth and secondary compound investments in the epiphytic lichensLobaria pulmonariaandHypogymnia occidentalistransplanted along an altitudinal gradient in British Columbia

We investigated altitudinal variation (550–1650 m) in relative growth rates (RGR) and carbon-based secondary compounds (CBSC) in the cephalolichen Lobaria pulmonaria (L.) Hoffm. and the chlorolichen Hypogymnia occidentalis L. H. Pike transplanted for 14 months in a U-shaped valley in inland southern British Columbia. Prior to transplantation, half of the thalli were treated with phosphorus (P) to examine effects of P on carbon allocation. Growth in L. pulmonaria was substantially higher in the toe-slope position, corresponding to much higher bark pH. Sixty-four percent of the variation in RGR was accounted for by positive pH effects and adverse effects of direct light exposure in the best subset multiple regression model. For H. occidentalis, 57% of the variation in RGR was accounted for by positive and negative effects of indirect and direct light, respectively. Neither altitude nor P had a noticeable effect on RGR, the former possibly reflecting a trade-off between orographic precipitation and inversion-boosted nocturnal dew in valley bottom localities. Neither was there any correlation between altitude and CBSCs, although treatment with P did significantly trigger secondary metabolism in L. pulmonaria, but not in H. occidentalis. No significant intraspecific relationship between growth and CBSC investments was noted.

Seasonal and spatial variation in carbon based secondary compounds in green algal and cyanobacterial members of the epiphytic lichen genus Lobaria

Acetone-extractable carbon based secondary compounds (CBSCs) were quantified in two epiphytic lichens to study possible effects of external factors (season and aspect) on secondary chemistry and to relate defense investments to biomass growth and changes in specific thallus mass (STM). At the end of four separate annual cycles starting in each of the four seasons, the cyanolichen Lobaria scrobiculata and the cephalolichen Lobaria pulmonaria (green algae as the primary photobiont and with localized Nostoc in internal cephalodia) were monitored in their natural forest habitats and after being transplanted at three contrasting aspects in open sites. Season strongly influenced most CBSCs. Medullary CBSCs in both species were twice as high in summer as in winter. Aspect hardly affected major CBSCs, whereas transplantation from forest to clear-cut slightly reduced these compounds. No major CBSCs in any species showed a trade-off with growth rate. Dry matter- as well as thallus area-based medullary CBSC contents increased with STM. The cortical usnic acid strongly increased with growth rate and followed spatial, but not seasonal variations in light exposure. Maximal CBSC levels during seasons with most herbivores is consistent with the hypothesis inferring that herbivory is a major selective force for CBSCs. Lack of trade-off between growth and defence investments suggests that these two processes do not compete for photosynthates.

The impact of secondary compounds and functional characteristics on lichen palatability and decomposition

Summary There has been much recent interest in understanding how functional traits of vascular plant species drive ecological processes such as herbivory and litter decomposition. In plants, these two processes are often driven by the same or similar suites of traits and therefore correlate across species. However, few studies have considered how traits of plant‐like life forms such as lichens determine species differences in their effects on ecological processes. This is despite the significant contribution of lichens to carbon and nutrient cycling in many environments. We collected 28 lichen species that differed in their growth form, substrate type and capacity to fix N, and determined key traits for each species. For each species, we performed a feeding bioassay using the generalist snail Cepaea hortensis and carried out a laboratory bioassay to assess decomposability. We did these tests both with intact lichen material containing natural concentrations of carbon‐based secondary compounds (CBSCs), and material that had been acetone rinsed to reduce concentrations of CBSCs, to evaluate the effect of CBSC on palatability and decomposability. We found that reducing CBSC concentrations greatly increased palatability for 17 species, and decomposability of 10 species. However, decomposability was correlated with several lichen traits while palatability was not, regardless of whether or not CBSCs were removed, and we therefore found no relationship between decomposability and palatability across species. Decomposability and palatability both varied, but in contrasting directions, among N‐fixing vs. non‐fixing lichens, lichens with different growth forms and those from contrasting substrate types. As such, N‐fixing lichens had higher decomposition rates but lower consumption rates than non‐fixing lichens, while foliose species had higher decomposition rates but lower consumption rates than fruticose species. Synthesis: We have shown that lichen CBSCs regulate key processes such as lichenivory and decomposition, that lichen decomposability but not palatability are related to traits, and that these two processes are unrelated across species. These results highlight the potential role of lichen species differences in influencing ecosystem processes relating to decomposition and nutrient cycling and the role that grazers may play in driving this.

Lichen Specific Thallus Mass and Secondary Compounds Change across a Retrogressive Fire-Driven Chronosequence

In the long-term absence of major disturbances ecosystems enter a state of retrogression, which involves declining soil fertility and consequently a reduction in decomposition rates. Recent studies have looked at how plant traits such as specific leaf mass and amounts of secondary compounds respond to declining soil fertility during retrogression, but there are no comparable studies for lichen traits despite increasing recognition of the role that lichens can play in ecosystem processes. We studied a group of 30 forested islands in northern Sweden differing greatly in fire history, and collectively representing a retrogressive chronosequence, spanning 5000 years. We used this system to explore how specific thallus mass (STM) and carbon based secondary compounds (CBSCs) change in three common epiphytic lichen species (Hypogymnia phsyodes, Melanohalea olivacea and Parmelia sulcata) as soil fertility declines during this retrogression. We found that STMs of lichens increased sharply during retrogression, and for all species soil N to P ratio (which increased during retrogression) was a strong predictor of STM. When expressed per unit area, medullary CBSCs in all species and cortical CBSCs in P. sulcata increased during retrogression. Meanwhile, when expressed per unit mass, only cortical CBSCs in H. physodes responded to retrogression, and in the opposite direction. Given that lichen functional traits are likely to be important in driving ecological processes that drive nutrient and carbon cycling in the way that plant functional traits are, the changes that they undergo during retrogression could potentially be significant for the functioning of the ecosystem.