Growth of reindeer lichens and effects of reindeer grazing on ground cover vegetation in a Scots pine forest and a subarctic heathland in Finnish Lapland

Reindeer lichens are an important component of northern ecosystems. The aim of this study was to measure the growth rate of terricolous lichens as it is a key parameter involved in productivity of these ecosystems and an important part of lichen tolerance to reindeer grazing. Furthermore, the natural succession and the long-term effects of reindeer grazing on lichen community characteristics in two contrasting habitats were investigated as well as the interactions between lichen cover and mosses and vascular plants. Biomass and coverage measurements were conducted in a lichen woodland and in a subarctic heath with grazed and ungrazed areas in northern Finland. Measurements spanning over 13 yr of undisturbed development show that the growth rate of Cladina stellaris can be as high as >0.17 g g⁻¹ produced annually, although in average growth rates were much lower. During the succession of ground vegetation, C. stellaris , C. rangiferina , C. mitis and Cetraria nivalis increased in biomass in fenced areas and were reduced most in biomass by reindeer in unprotected areas. Reindeer grazing and trampling seem to change the vegetation towards a type that is dominated by small dwarf shrubs, bare soil and minute-cup lichens ( Cladonia spp.). Removing the lichen layer by reindeer may reduce natural regeneration of pine trees as implied by increasing numbers of pine seedlings with increasing lichen cover.     

Cladonia lichens as the most effective and essential pioneers in strongly contaminated slag dumps

The epigeic lichen biota of ten post-smelting slag dumps were studied to identify the ecological importance of Cladonia species in spontaneous succession within artificial areas. Species diversity, abundance, constancy and cover indices were determined at the initial stages of vegetation establishment. Substrates under lichen vegetation were characterised in terms of the pH, organic C and total N content, C/N ratio and Zn, Pb, Cd and As concentration. Species occurrence and abundance indices were related to habitat type, enabling the ability to identify species-specific responses to environmental factors. Despite the unfavourable and artificial habitat conditions at dump sites, some Cladonia appear abundant and form swards, with Cladonia rei as a dominant species. They appear to be well-adapted for slag substrate and their special arrangement illustrative of site contamination. Certain sward-forming Cladonia lichens play a fundamental role as effective pioneers in the initial stage of succession and provide stable plant cover development.     

Antimicrobial activity of extracts of the lichens <Emphasis Type="Italic">Cladonia furcata,Parmelia caperata,Parmelia pertusa,Hypogymnia physodes</Emphasis> and <Emphasis Type="Italic">Umbilicaria polyphylla</Emphasis>

Antimicrobial features of acetone, methanol and aqueous extracts of lichens of Cladonia furcata, Parmelia caperata, Parmelia pertusa, Hypogymnia physodes and Umbilicaria polyphylla were investigated by two different methods at the same time. Testing of antimicrobial activities of extracts from five species of lichens was performed by disc diffusion test in relation to Gram-positive and Gram-negative bacteria and fungal organisms, and through determination of minimal inhibitory concentration (MIC) by Broth Tube Dilution method. The obtained results indicated that acetone and methanol extracts of all investigated lichens in different concentrations manifested selective antibacterial and antifungal activity. That activity was more evident in relation to Gram-positive, than Gram-negative bacteria and fungal organisms. Acetone and methanol extracts of lichens Parmelia pertusa, Hypogymnia physodes and Umbilicaria polyphylla inhibited the growth of all tested microorganisms, most of all of lichens Cladonia furcata and Parmelia caperata. Although, the methanol extracts were generally the most active against the test organisms, the lowest MIC value was measured for acetone extract of species Cladonia furcata 0.39 mg/mL in relation to bacterium Bacillus subtilis. Aqueous extracts of investigated lichens were inactive against all tested organisms.     

Lecanoric acid,a secondary lichen substance with antioxidant properties from <Emphasis Type="Italic">Umbilicaria antarctica</Emphasis> in maritime Antarctica (King George Island)

Eight lichen species, Cetraria aculeata, Cladonia furcata, Pseudephebe pubescens, Sphaerophorus globosus, Stereocaulon alpinum, Umbilicaria antarctica, Usnea antarctica and Usnea aurantiacoatra, were collected from King George Island, maritime Antarctica, for the evaluation of antioxidant activities. Anti-linoleic acid peroxidation activity, free radical scavenging activity, reducing power and superoxide anion scavenging activity were assessed of methanol and acetone extract of the lichens in vitro. Extract of Umbilicaria antarctica, Cladonia furcata, Sphaerophorus globosus and Usnea antarctica were found to have strong in vitro antioxidant properties. In general, acetone extract exhibited stronger activities than methanol extract. The activity-guided bioautographic TLC and HPLC analysis demonstrated that lecanoric acid was the main antioxidant compound in the acetone extract of Umbilicaria antarctica, the most potent antioxidant lichen species among the test species. The results suggested that several Antarctic lichens and their substances can be used as novel bioresources of natural antioxidants.     

Zn/Pb-tolerant lichens with higher content of secondary metabolites produce less phytochelatins than specimens living in unpolluted habitats

Many lichens can cope with heavy-metal stress, however, the mechanisms of lichen tolerance are still not fully understood. Some lichen secondary metabolites (depsides and depsidones), produced in lichens by the fungal symbiont and accumulated on the outer surface of its hyphae, are supposed to play an important role in the extracellular immoblilization of heavy metals. Lichen photobionts (algal partners in the symbiosis), although surrounded by the mycobiont hyphae, may also accumulate high amounts of trace metals. This can lead to physiological disruptions and morphological damage in algal cells and hence affect the lichen physiological status. We hypothesized that lichen species/specimens living in heavily polluted sites and showing HM tolerance possess a higher content of secondary metabolites than those living in unpolluted sites. Hence, their photobionts can be better protected from the excess of metal ions and need to produce less metal-complexing phytochelatins (PCn) to combat metal toxicity. Specimens of Hypocenomyce scalaris, Cladonia furcata and Lepraria spp. sampled from Zn/Pb-polluted and control sites were compared for the accumulation of Zn/Pb and secondary metabolites, as well as for their production of phytochelatins and glutathione in response to experimental Zn or Pb exposure. Generally, the lichen specimens sampled from the HM-polluted site contained higher amounts of Zn and Pb as well as lichen substances (different depsides and depsidones) than those from the control site. A strong positive correlation was found between the accumulation of secondary metabolites and Zn/Pb accumulation (R² = 0.98 and 0.63, respectively). For the first time, production of phytochelatins (PC_2-3) in response to Zn and Pb (50-200 μM) exposure was found in H. scalaris, L. elobata, L. incana and C. furcata. In both species of Lepraria also cysteine, a substrate for GSH and PCs synthesis was detected. The lichens from the polluted site produced under the same exposure conditions, or in response to higher metal concentrations, lower amounts of PCn than those sampled from the control site. It strongly suggests that less Zn and Pb ions reached the photobiont cells of the lichens containing higher amounts of secondary metabolites (lecanoric, fumarprotocetraric, stictic, constictic acids, antranorin). The results obtained support the putative role of some metabolites in heavy-metal tolerance of the lichens inhabiting metal-polluted habitats.     

The effect of the substrate on the growth and reproduction of the lichens Cladonia rangiferina and C. mitis

The growth rate and percentage of fertile individuals were studied in the lichens Cladonia rangiferina and C. mitis growing on various substrates in a heath-lichen pine forest and a bilberry-cowberry pine forest with Pleurozium and other mosses, both on the White Sea coast. The substrates were soil, rocks, and pine wood. The proportion of different forms of C. rangiferina growing on the same substrate was compared between the two forest types. C. rangiferina f. rangiferina was more abundant on soil, while C. rangiferina f. crispata dominated on wood. The effect of the substrate on the development of different lichen forms is discussed. The maximum growth rate and the maximum percentage of fertile individuals were recorded in the lichens growing on soil; these parameters were minimal in the lichens growing on wood. These differences were more pronounced in C. rangiferina than in C. mitis. The effects of the nutritive quality and the humidity of the substrate on the growth rate and apothecia formation in lichens are discussed.     

Invasiveness of Campylopus introflexus in drift sands depends on nitrogen deposition and soil organic matter

Question: Does the neophyte moss Campylopus introflexus invade more often in drift sand pioneer vegetations under high nitrogen (N) deposition? Location: Fourteen inland dune reserves in The Netherlands over a gradient of atmospheric N deposition. Methods: A transect study, dispersal experiment and culture experiment were carried out. In the transect study, the establishment of C. introflexus and lichens was measured in pure mats of Polytrichum piliferum, an early succession stage. The overall presence of C. introflexus in the area was also estimated. In the dispersal experiment, fragments of C. introflexus and lichen species were sown in P. piliferum mats at two sites with high and low N deposition. In the culture experiment C. introflexus fragments were grown on soil with different carbon (C) content and N dose. Results: The Campylopus:lichen ratio was positively correlated (r²=0.61) with the atmospheric ammonia concentration. Campylopus began to dominate at an ammonia air concentration of 7 μg m^?3, correlated with the overall presence of the species in the sites investigated. Survival of sown Campylopus fragments was significantly higher and the endangered lichen Cladonia strepsilis significantly lower in the site with a high ammonia concentration. Survival of Cladonia coccifera and Cladonia portentosa was high at both sites. Experimental growing of C. introflexus showed significant responses to both C content and N, although C content showed the strongest effect. Conclusions: Growth of C. introflexus is affected by soil C content and N deposition. Inland dune reserves under high N deposition risk loss of lichen‐dominated vegetations because of moss encroachment.     

Growth and nitrogen relations in the mat-forming lichens Stereocaulon paschale and Cladonia stellaris

BACKGROUND AND AIMS: Mat-forming lichens in the genera Stereocaulon and Cladonia have ecosystem-level effects in northern boreal forests. Yet the factors affecting the productivity of mat-forming lichens are not known. The aim of the presented work was to investigate whether mat-forming lichens adapted to low N availability employ N-conserving mechanisms similar to those of vascular plants in nutrient-poor ecosystems. Specifically, the following questions were asked: (a) Do lichens translocate N from basal areas to apical growth areas? (b) Are the quantities of N translocated of ecological significance. (c) Is lichen growth dependent on tissue N concentration [N]. METHODS: Two different, but complementary, field experiments were conducted using the mat-forming N2-fixing Stereocaulon paschale and non-fixing Cladonia stellaris as model species. First, N translocation was investigated by feeding lichens with Na(15)NO3 either directly to the apex (theoretical sink) or to the basal part (theoretical source) and observing the redistribution of (15)N after a growth period. Secondly, growth and variation in [N] in thalli of different lengths was measured after a growth period. KEY RESULTS: (15)N fed to lower parts of lichen was translocated towards the growing top, but not vice versa, indicating physiologically dependent translocation that follows a sink-source relationship. In the growth experiment where thalli were cut to different lengths, the significant decrease in [N] in apices of short vs. longer thalli after a growth period is consistent with internal relocation as an ecologically important source of N. CONCLUSIONS: The presented results demonstrate that internal recycling of N occurs in both species investigated and may be ecologically important in these mat-forming lichens under field conditions. The higher nitrogen use efficiency and relative growth rate in C. stellaris in comparison with S. paschale probably enable C. stellaris to dominate the ground cover vegetation in dry boreal coniferous forests under undisturbed conditions.     

Removal of secondary compounds increases invertebrate abundance in lichens

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.     

地衣型真菌次级代谢产物抗菌活性初步研究

地衣是一种传统的民族药物,能产生多种具有活性的物质。该研究对地衣型真菌(Xanthoria elegans,Myelochroa indica,Ramalina peruviana,Cladonia macilenta,Nephromopsis pallescens,Cladonia coccifera)进行液体培养,2个月后,培养液用乙酸乙酯萃取后获得初提物。该研究采用抑菌圈法评价地衣型真菌初提物对7种致病细菌(Bacillus subtilis,Bacillus cereus,Vibrio parahaemolyticus,Straphylococcus haemolyticus,Pseudomonas aeruginosa,Staphylococcus aureus,Micrococcus luteus)的抗菌活性,并测定最低抑菌浓度(MIC)。结果表明:6种地衣型真菌的初提物均具有一定的抗菌活性,且不同培养基对地衣型真菌产生抗菌物质有显著影响。其中,R.peruviana在MY液体培养基中所产生的次级代谢产物对金黄色葡萄球菌、藤黄微球菌、溶血性葡萄球菌、铜尿假单胞菌具有抑制效果,但在YMG培养基中所得初提物对供试7种致病细菌不具有抑菌效果。X.elegans在YMG培养基中所得初提物对枯草芽孢杆菌具有明显抗菌活性,其抑菌圈直径可达17.77 mm。该研究证实不同地衣型真菌液体培养初提物具有抗菌活性,不同的培养基也直接影响地衣型真菌抗菌效果。该研究结果为地衣型真菌的进一步研究及民族药的开发利用奠定了基础。     

Analysis of the distribution of epiphytic lichens within homogeneousFagus sylvatica stands along an altitudinal gradient (Mount Olympos,Greece)

Epiphytic lichen vegetation onFagus sylvatica sas studied in 4 sites along an altitudinal gradient from 930 to 1500 m on SE facing slopes of Mount Olympos (Greece). The crucial factor determining the spatial heterogeneity of epiphytic lichens onF. sylvatica is the altitude and not the height on the trunk at which lichen community is established. 17 out of 26 taxa are confined to a particular elevation range, while another three are clearly ubiquitous in their distribution. The number of lichen species at breast height is higher than at the base of the trunks. The results were compared with those gathered earlier in an analogous study on the vertical distribution of epiphytic lichens onPinus nigra along an altitudinal gradient from 750 to 1510 m of the same mountain. Comparison suggests that spatial heterogeneity of epiphytic lichens onF. sylvatica is different from the one onP. nigra.     

新疆天山西部山脉森林生态系统地衣群落结构的初步研究

 根据多年的研究资料并采用聚类分析方法对分布在天山西部山脉的地衣群落结构进行了研究。结果表明,分布在该地区的地衣种类形成了6种群落:1) 土星猫耳衣（Leptogium saturninum)＋矮石蕊(Cladonia humilis)＋兰灰蜈蚣衣（Physcia caesia）群落;2）黑穗石蕊（Cladonia amaurocr     

Iron and phosphate uptake explains the calcifuge–calcicole behavior of the terricolous lichens Cladonia furcata subsp. furcata and C. rangiformis

Mechanisms causing the calcifuge–calcicole behavior of lichens are largely unexplored. Studying the case examples of two closely related terricolous lichens, the calcifuge Cladonia furcata subsp. furcata and the calcicole C. rangiformis, we found that preference for acidic or calcareous soils in these lichens is related to iron and phosphate uptake as in vascular plants. In laboratory studies, the calcicole species was more efficient in the intracellular uptake of Fe³⁺ and phosphate at pH 8 than the calcifuge species. At pH 3, intracellular uptake of Fe²⁺ in the calcicole species significantly exceeded that in the calcifuge species suggesting that calcicole lichens suffer from toxicity symptoms by excess Fe²⁺ at acidic sites. Though these observations parallel findings from calcifuge and calcicole vascular plants, mechanisms leading to the different iron and phosphate uptake characteristics in the studied calcifuge and calcicole lichens may differ from those in vascular plants and should be the topic of future research. A role of the depside atranorin in facilitating iron uptake by reducing Fe³⁺ in the apoplast is hypothesized.     

STRUCTURAL STUDIES ON THE GALACTOMANNANS OF LICHENS OF THE GENUSCLADONIA

Galactomannans were isolated fromCladonia signata,C. furcata,C. imperialis, andC. clathratavia successive alkaline extraction and precipitation with Fehling solution and Cetavlon. They were investigated using¹³C-NMR spectroscopy, methylation analysis, and Smith degradation, and their specific rotations and monosaccharide compositions determined. As with galactomannans of otherCladoniaspecies, they contained (1→6)-linked main chains of α-mannopyranose, which were non-substituted (structure4in Fig. 2), mono-substituted at O-2 with α-mannopyranose (structure6) or α-galactopyranose (structure1), O-4 with β-galactopyranose (structure2), and disubstituted at O-2 and O-4 with α-mannopyranosyl and β-galactopyranosyl units, respectively (structure5). Disubstitution was present to a greater extent in the galactomannans ofC. clathrataandC. imperialisthan in those ofC. signataandC. furcata. In the case of the galactomannans ofC. furcata,C. clathrata, andC. imperialis, substitution also occurred at O-2 withO-β-galactofuranosyl-(1→6)-O-α-mannopyranosyl units (structure7). As observed in previous investigations, the C-1 portion of the¹³C-NMR of mannose-containing polysaccharides is typical of the lichen species. However, those of galactomannans ofC. imperialisandC. clathrataare almost identical and, although other chemical data showed many structures in common, some differences were evident.     

Nutritive value of terricolous lichens for reindeer in winter

Abstract:In contrast to virtually all other species of ruminants, reindeer (Rangifer tarandus) eat substantial amounts of lichens in winter. Several different species of lichens are eaten and most are highly palatable. The chemical composition and the in vitro digestibility of six species of terricolous lichens commonly eaten by reindeer were measured. Ruminal inoculum was obtained in winter from six free-ranging reindeer that had been grazing on natural pastures in northern Norway. Analysis of the plant parts recovered from their rumens confirmed that the animals had been eating a mixed diet that included both lichens and vascular plants. The chemical composition and the in vitro digestibility of the different species of lichens varied considerably between species and genera. Cetraria islandica, C. nivalis andCladonia arbuscula were highly digestible [69–77% dry matter (DM)], Stereocaulon paschale was poorly digestible (43% DM), whereas Cladonia stellaris and C. gracilis had an intermediate digestibility (56–57% DM). Mixing Cladonia stellaris with vascular plants (50:50) had no effect on the in vitro digestibility of the combined substrates. The in vitro digestibility of Cladonia stellaris in inoculum from two captive reindeer which had had no access to lichens was very low (10% DM). This shows that the source of inoculum used for digestibility trials has a major effect on the apparent digestibility of the substrates. Evidently, the extent to which reindeer are able to utilize lichens depends on the species that are selected and on what the animals have been eating recently.     

Lichens at Dungeness

LAUNDON, J. R., 1989. Lichens at Dungeness . The lichen vegetation at Dungeness is reviewed. The most important lichen communities are of three types. Firstly, there are the crustose lichens which colonize the bare pebbles. Secondly, the tracts of rich Cladonia turf which form the lichen heath and which play a major role in the vegetation cycle. Thirdly, there are the epiphytes on Prunus scrub. In addition, lichens on artificial substrates, especially wooden fences, are noteworthy. About 150 species occur, amongst the most interesting being Cladonia mitis, Lecanora helicopis, Lecidea insidiosa, Parmelia soredians, Rinodina aspersa and Umea glabrata. A new combination is made in the genus Porpidia. The term 'bird farming' is published. The fragmentation of the vegetation by building and gravel extraction receives critical comment.     

Estimating plant abundance using inflated beta distributions: Applied learnings from a lichen–caribou ecosystem

Quantifying abundance and distribution of plant species can be difficult because data are often inflated with zero values due to rarity or absence from many ecosystems. Terrestrial fruticose lichens (Cladonia and Cetraria spp.) occupy a narrow ecological niche and have been linked to the diets of declining caribou and reindeer populations (Rangifer tarandus) across their global distribution, and conditions related to their abundance and distribution are not well understood. We attempted to measure effects related to the occupancy and abundance of terrestrial fruticose lichens by sampling and simultaneously modeling two discrete conditions: absence and abundance. We sampled the proportion cover of terrestrial lichens at 438 vegetation plots, including 98 plots having zero lichens. A zero‐inflated beta regression model was employed to simultaneously estimate both the absence and the proportion cover of terrestrial fruticose lichens using fine resolution satellite imagery and light detection and ranging (LiDAR) derived covariates. The probability of lichen absence significantly increased with shallower groundwater, taller vegetation, and increased Sphagnum moss cover. Vegetation productivity, Sphagnum moss cover, and seasonal changes in photosynthetic capacity were negatively related to the abundances of terrestrial lichens. Inflated beta regression reliably estimated the abundance of terrestrial lichens (R² = .74) which was interpolated on a map at fine resolution across a caribou range to support ecological conservation and reclamation. Results demonstrate that sampling for and simultaneously estimating both occupancy and abundance offer a powerful approach to improve statistical estimation and expand ecological inference in an applied setting. Learnings are broadly applicable to studying species that are rare, occupy narrow niches, or where the response variable is a proportion value containing zero or one, which is typical of vegetation cover data.     

Transplantation of lichen thalli: a case study on Cetraria islandica for conservation and pharmaceutical purposes

Changes in biomass growth and chemical composition of transplanted Cetraria islandica lichen thalli were investigated in a natural stand in the Bory Tucholskie region (northern Poland) over 3 yr. Transplants consisted of either 3.85 or 7.88 g fragments, 12.62 g clumps, or control plots where all vegetation and lichens were removed. The initial mass of C. islandica significantly influenced the percentage cover of investigated lichens at the study plots. At the last set of study measurements the highest cover, biomass accumulation (4×) and mean final mass (31.99 g m⁻²) of C. islandica were recorded at the study plots at which 7.77 g fragments were transplanted. Analysis showed that the content of the determined chemical compounds in each sample were similar. The potential influence of other species growing at the study area on the occurrence of C. islandica was also examined through monitoring species diversity in the cleared forest floor plots. Altogether 18 species of plants and lichens were recorded in study plots, and among most abundant species Dicranum polysetum and Pleurozium schreberi were observed. The mean percentage cover for C. islandica was 14.61 %.     

Relations between understorey vegetation,site productivity,and environmental factors in Pinus sylvestris L. stands in southern Finland

Relations between understorey vegetation, site productivity, and environmental factors in mature stands of Pinus sylvestris L. in southern Finland were studied using data from 234 sample plots. The (weighted) Pearson product-moment correlation between site index and the first ordination axis of detrended correspondence analysis (DCA) was 0.83. The correlation was clearly better where site index was above 21 m (r=0.83) than where it was less (r=0.33). Partial canonical correspondence analysis (CCA) indicated that the residual compositional variation, not related to site fertility as expressed by site index, can be partly ascribed to high moisture regime with poor drainage (paludification), soil deposit type, soil stoniness, and macroclimate, and in part to unexplained sources. The unique influence of canopy cover on the vegetation composition was non-significant (p<0.05) and that of stand age too small to be detected in the vegetation data. The results support Cajander's forest site type theory according to which there exists a dominant relationship between understorey vegetation and site productivity in boreal forest ecosystems. Cladonia sectio Cladonia lichens, exluding Cladonia uncialis, were treated collectively, and abbreviated as Cladonia spp. The most frequent of these species were C. gracilis, C. cornuta, C. fimbriata, C. deformis, C. cenotea, C. botrytes, C. crispata, C. pyxidata, C. cornuta, and C. furcata.     

Antioxidant and antimicrobial activity of some lichen species

The aim of the research is to explore the overall in vitro antioxidant activity, total phenol content, reduction power and antimicrobial activity of the methanol extracts of the lichens Cetraria pinastri, Cladonia digitata, Cladonia fimbriata, Fulgensia fulgens, Ochrolechia parella and Parmelia crinita. The methanol extract of the Cetraria pinastri showed a strong antioxidant activity, whereas the extracts of the species Fulgenesi fulgens, Cladonia fimbriata and Parmelia crinita showed the moderate one and the extract of the species Ochrolechia parella and Cladonia digitata the weak one. The methanol extract of the lichen Cetraria pinastri had the biggest total phenol content (32.9 mg/g of the dry extract). A certain correlation was established between the antioxidant activity and the total phenol content for the researched lichen extracts. The work also explores the antimicrobial activity of the methanol extracts of the mentioned species of lichens against six bacterial and eleven fungi species by the disc-diffusion method and by establishing the minimum inhibitory concentration (MIC). The methanol extracts of the lichens Cetraria pinastri and Parmelia crinita showed the strongest both antibacterial and antifungal activity against most of the tested microorganisms. These researches suggest that the lichens Cetraria prunastri can be used as new sources of the natural antioxidants and the substances with antimicrobial features.