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Coastal Lichens Distribution Patterns and Adaptive Strategies

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DOI: 10.1007/978-3-030-17854-3_34-1

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Coastal Lichens
Distribution Patterns and Adaptive Strategies

Angella V. Sonina and Vera I. Androsova

Contents
1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Freshwater Lichens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3 Marine Lichens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
4 Adaptations of Marine Lichens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
5 Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

Abstract
Lichens are a symbiotic complex of autotrophic (algae, Cyanobacteria) and
heterotrophic (fungi) components that have developed during evolution in coastal
ecosystems in the process of adaptation of algae and fungi to terrestrial habitats.
Lichens are highly adapted to extreme habitats including the littoral (or intertidal)
zones of coasts. In this chapter, we present developmental stages of aquatic lichen
investigations: freshwater and marine lichens. The issues of species diversity of
coastal lichens, their ecology, and adaptations to the coastal marine environment
are described. The leading factors affected the epilithic lichen cover of coasts, and
freshwater habitats are at a distance from the waterline and substrate character-
istics. Substrate characteristics, especially near the waterline, depend on the wave
rhythm. On the coasts of freshwater bodies, four zones are recognized based on
flooding duration and lichen ecology. Lichen zones of fresh and marine coasts are
distinguished by their species composition: on sea coasts halophytes are predom-
inant and on freshwater shores – hydrophilic lichens. Marker species of lichens
were identified for each zone. For the littoral zone, the intrazonal structure of
lichen flora was shown. In the adaptation of symbiotic organisms, such as lichens,
both symbionts take part: mycobiont and photobiont. Morphological and struc-
tural adaptations are mainly associated with mycobiont variability: the presence

A. V. Sonina (*) · V. I. Androsova

Petrozavodsk State University, Petrozavodsk, Republic of Karelia, Russia

© Springer Nature Switzerland AG 2020 1

M. N. Grigore (ed.), Handbook of Halophytes,
https://doi.org/10.1007/978-3-030-17854-3_34-1
2 A. V. Sonina and V. I. Androsova

of morphotypes, structural features of the reproductive organs, and anatomical

layers. Photobiont is responsible for functional adaptations: the variability of the
amount of photosynthetic pigments and the synthesis of various substances that
ensure the resistance of lichen to salt stress. The photobiont provides synthesis of
osmolytes, and these process patterns can change depending on the photobiont
species (strain), as well as for one photobiont in different coastal conditions. The
distribution of epilithic lichen species on the coasts is therefore also ensured by
the functions of the photobiont. The important component of the lichen associ-
ation is the microbial complex; however its role in adapting lichens to coastal
environment is still not clear. The perspective of studying coastal lichens is
determined by a rather poor knowledge of species diversity and ecology of this
group, including physiology, biochemistry, and genetics based on modern
research methods.

Keywords
Coastal lichens · Epilithic lichens · Freshwater lichens · Marine lichens · Tidal
zone · Adaptation

1 Introduction

Lichens are a symbiotic complex of autotrophic (algae, Cyanobacteria) and hetero-

trophic (fungi) components that have developed during evolution in coastal ecosys-
tems in the process of adaptation of algae and fungi to terrestrial habitats. Lichens in
the current literature are considered as a result of the symbiogenesis between fungi
and algae during evolution and occupation of terrestrial environment by eukaryotes
as well as strategic way of adaptation of fungal organisms. Therefore, mycologists
consider lichens as lichenized fungi and classified them in the system of the Perfect
fungi (Hawksworth and Honegger 1994).
In fact, lichenology is a relatively new branch of biological science. Currently,
along with the study of such traditional issues as the diversity of lichen species, the
ecology and physiology of these complex organisms, more and more are being
studied.
In ecosystems, lichens play the role of primary producers and contribute to
organic matter formation and nitrogen cycling. Due to their high resistance, lichens
are able to survive in places that most other organisms are unable to occupy such as
dry, cold deserts, tundra communities, rocky habitats, etc. Lichens play a significant
ecological role as the primary colonizers (frozen lava flows, coastal cliffs, glaciers,
etc.) and participate in mineral weathering and pedogenesis. Meanwhile lichens are
characterized by poor competitive ability in relation to higher plants (Tarasova et al.
2012). Lichens can colonize habitats with significant differences in water ability.
Nevertheless, it was noted that freshwater and marine lichen-forming fungi are rarely
studied by lichenologists and mycologists (Hawksworth 2000).
Coastal Lichens 3

Species diversity and ecological features of freshwater and marine lichens

(study history). Specific conditions for the biota development are formed in fresh-
water and sea coastal habitats. The coastal line presents the contact zone between
aquatic and terrestrial ecosystems, which penetrate into each other and form habitats
different from initial environments. The spatial organization of coastal communities
is associated with the influence, on the one hand, of aquatic ecosystems and, on the
other, of terrestrial ones (Markovskaya et al. 2010; Sonina 2014).
Since halophyte plants have been extensively studied that it is associated with
global problems of humanity due to predicted salt stress linked to climate change
(Hasegawa et al. 2000 etc.), marine lichens have been specifically studied more
recently. However, a number of ecological and floristic studies have emphasized the
freshwater lichens. Freshwater and marine lichens are a represented group of aquatic
lichens. Both marine and freshwater lichens are facing strong physical impact on all
surface structures due to the impact of the water current. Vertical zonations of lichens
are characteristic of marine and freshwater habitats. Often, researchers reported
results of comparative study, and this obliges us to consider both of these groups.
The study of aquatic lichens can be divided into three periods using the chrono-
logical and the thematic approaches.
The first period is characterized by the concentration of floristic studies in the
nineteenth century. In Europe, aquatic lichens are first mentioned in the works of
Acharius (1798–1810), Lindsay (1856), and Arnold (1868–1887) (cited by Savich
1950). Meanwhile, along with lists of lichens, some ecological descriptions of
aquatic species and their habitats preferences were reported. In 1856 W.L. Lindsay
described the ecological features of Dermatocarpon miniatum var. complicatum
found on flooded boulders along the Tai River.
In several works of F. Arnold in 1868–1887 (cited by Savich 1950), the species
composition of aquatic lichens of streams and mountain lakes is discussed. Moreover,
the author classified water lichens depending on habitat conditions, with different
levels of flooding. The researcher recognized three groups of species: the first group
includes lichens that constantly grow in water; the second group, lichens which can
grow both in water and outside water; and the third group, lichens which are resistance
to short-term flooding (cited by Savich 1950). At the same period of time, the term
“hydrophilic lichens” began to be used (Tsukal 1896: cited by Savich 1950).
The second stage included ecological and floristic research of the twentieth
century. In an earlier study, A.A. Elenkin (1921) reported the species Verrucaria
aethiobola Wahlenb. from underwater habitat. (¼ V. aquilella Nyl.) (in the Moscow
region). Later, L. G. Ramensky (1922) mentioned the species Collema ramenskii
Elenk. from Lake Ukshezero (Republic of Karelia) at a depth of 1.5 m (cited by
Savich 1950). A classical paper of Savich (1924) about water lichens of Karelia and
Kamchatka included a report of findings of hydrophilic lichen species: Verrucaria
hydrela Ach., V. aethiobola Wahlenb., V. aquatilis Mudd., Bacidia imendata Körb.,
and Aspicilia aquatica Körb. (cited by Savich 1950).
Many studies demonstrated that the group of aquatic lichens is not homogeneous
and the species differ in habitat preferences depending on the flooding regime (Smith
1921). The research on ecology of aquatic lichens by Savich (1950) has suggested that
4 A. V. Sonina and V. I. Androsova

species are associated with various conditions of flooding regime: (1) lichens that
occur mostly in underwater substrates or in places periodically flooded with water; (2)
lichens that use moisture from splashing waterfalls, surfs, etc.; and (3) lichens that can
withstand prolonged accidental flooding, while they usually occur outside the water
level. A similar extensive study was carried out by Räsänen (1927) in the western part
of Finland. Savich (1950) and Räsänen (1927) made the conclusion about the funda-
mental differences between the freshwater and coastal lichen communities. The
researches also recognized two groups of lichens on stones irrigated with freshwater
and salt water. Räsänen (1927) characterized lichens on underwater stones as stable
organisms to constant influence of water level. He described the lichen’s ability as “the
most biologically resistant” to such environmental conditions.
In this period adaptations of aquatic lichens are reported in some studies. A. L.
Smith (1921) described adaptive morphological characteristics of this group of
lichens such as crustose type of thallus and presence of perithecia.
Special attention of researchers to the study of freshwater and marine lichens was
formed by the middle of the twentieth century. By this time, about 700 species of
lichens and allied fungi were reported for tidal zones, and 200 were known – from
freshwater rivers and lake shores (Santesson 1939).
Based on observations and analysis of the distribution of lichens on the shores of
freshwater bodies and sea coasts, four lichen zones are distinguished (Scott 1967;
Santesson 1939) according to the flooding duration of lichens as well as on the
ecology of species.
At the third stage (modern period, the end of twentieth century, twenty-first
century) of studying aquatic lichens, great interest was brought to their ecology.
Many works focused on structure as well as the patterns and mechanisms of the
formation and functioning of lichen cover in aquatic environment. Studies on the
ecological and physiological adaptation mechanisms of aquatic lichens to conditions
with different levels of humidification and hydration of thalli were developing, and
data started to accumulate (Lange et al. 1986). Nevertheless, many fundamental
aspects of this group of lichens remain uncertain.
Most work on coastal and freshwater lichens has been carried out in temperate
regions (Hawksworth 2000). The main works are concentrated in the territories of
England, Northern Europe, and northwest and northeast of America (Pentecost
1977; Hale 1984; Giavarini 1990; Gilbert and Giavarini 1993; Gilbert 2003; Hansen
et al. 2006; Wirth et al. 2007). The works are related both to the coasts of freshwater
lichens, which is mainly determined by the increase in human attention to the sea
coasts and of freshwater shores.

2 Freshwater Lichens

Many current studies demonstrated new species findings of freshwater habitats. For
example, in 36 perennial springs of the eastern Italian Alps, 34 lichen species were
found including 1 new to Italy (Nascimbene and Nimis 2007). Verrucaria
elaeomelaena and V. funckii were suggested by authors as potential indicator species,
Coastal Lichens 5

on calcareous and siliceous springs, respectively. Among the 1642 species of lichens
known from Poland (Fałtynowicz and Kossowska 2016), 56 are freshwater lichens.
In a recent paper, seven freshwater lichen species newly recorded from the Western
Carpathians in Poland were reported (Matura et al. 2017). Sedelnikova for the
regions of the Altai-Sayan mountain region recognized a group of limestone lichens,
which are associated with periodic flooding (Sedelnikova 2001). She also described
a new species for science, Aspicilia popovae Sedeln., which occurs underwater
during summer (Kulikova et al. 2008). Macry (1990) indicates for the lichen flora
of the Baikal ridge several new species from the family Verrucariaceae, distributed
on the stones of rivers and streams and periodically flooded with water.
In total, the number of described freshwater lichens is about 250 species
(Hawksworth 2000). The highest percentage of aquatic lichen species belongs to
the Verrucariaceae family (more than 20 species of lichenized fungi) (Thüs et al.
2014). However, the complete species diversity of freshwater lichens has not yet
been identified, and further studies are required. In fact, there are practically no data
for subtropical and tropical regions, as well as for temperate regions in the Southern
Hemisphere (Thüs et al. 2014).
Numerous reports provide data on the vertical zonation patterns of aquatic lichens
which are related to different levels of flooding and characterized by specific species
lichen composition. For the first time, these data was represented in the work of R.
Santesson (1939), where lichen zones on lake shores in Sweden were characterized
in connection with the flooding duration of certain lichen species. The authors also
identified “lichen line” below which foliose lichens could not survive. Moreover,
analysis of the algal partner established that species with symbiont from the genus
Trebouxia are sensitive to flooding and unstable to it.
Comparing the lichen flora of two mountain streams in England, A. Pentecost
arranged the lichen species in ecological series depending on the water factor
(Pentecost 1977). The work of Gilbert and Giavarini (1997) demonstrated the
influence of the water acidity of lakes in England on the species composition of
epilithic lichens. Those researchers have also recognized four zones for studied
freshwater habitats with characteristics of lichen species composition (Gilbert and
Giavarini 1997):

1. Underwater zone: Collema dichotomum, Hymenelia lacustris, Placynthium

flabellosum, Poroscyphus kenmorensis, Staurothele jissa, Verrucaria aquatilis,
V. funckii, and V. praetermissa
2. Fluvial mesic zone: Aspicilia laevata, Bacidia inundata, Dermatocarpon
meiophyllum, Ephebe lanata, Porpidia hydrophila, Verrucaria aeithiobola, and
V. latebrosa
3. Fluvial xeric zone: Aspicilia caesiocinerea, Catillaria chalybeia, Massalongia
carnosa, Porina lectissima, and Porpidia soredizodes
4. Fluvial terrestrial zone: with numerous terrestrial species, including “lichen line
indicators” such as Parmelia conspersa
6 A. V. Sonina and V. I. Androsova

Fig. 1 The degree of surface

structure of the substrate at a
different distance from the
waterline (points: 1 smooth
substrate, graininess is not
expressed, 2 smooth grained
substrate, 3 finely grained
substrate, 4 medium-grained
substrate, 5 coarse-grained
substrate, 6 grained substrate
with cracks and chips; ***p
<0.001)

Similar zones are reported for other regions of Europe (Hawksworth 2000; Coste
2009). In the northwest of Russia, on the shores of rivers and the Onega Lake, four
zones are also recognized with species of the genus Verrucaria and Staurothele as
indicators of zones in relation to water level (Sonina et al. 2000). Based on multi-
dimensional statistics, the same authors showed that in conditions of freshwater
habitats, the leading factors which affected the epilithic lichen cover are at a distance
from the waterline and substrate characteristics (Sonina et al. 2000). In addition, the
correlation between these two habitat characteristics was revealed: with increasing
distance from waterline substrate, it becomes morphologically more homogeneous
(Fig. 1).
In 2002, Thüs published a monograph on the problematic group of freshwater
lichens “Hydroverrucaria” in Central Europe with a description of their ecology.
Аquatic lichens in France have been extensively studied by Coste (2009), and a
current classification was proposed for the phytosociology of epilithic coastal
lichens. The author proposes to combine freshwater lichens into one class of
hydrophilic lichens, among which three orders standout corresponding to the differ-
ent times of contact with water, six alliances that depend on bioclimatic stage, and
finally several associations by species composition.
Canadian researchers, based on the methods of multidimensional statistics, stud-
ied the structural organization of epilithic lichen sinusia of lacustrine sandstones of
the Canadian Rockies (John and Dale 1990) and the cliffs of Niagara Falls (Matthes
et al. 2000). In North America, in Idaho, Rosentreter (1984) indicated species of the
genus Verrucaria as indicators of low water levels and the species Dermatocarpon
reticulatum as characteristic of the normal flood zone. Similar zoning patterns have
been described in Africa around basins and water channels in granite mines in
Zimbabwe (Scott 1967). In some temperate regions, the species Parmelia conspersa
is considered as an indicator of the maximum height of floodwaters (Hale 1984).
Zones were related to the distribution in the upper zone of the yellow-green lichen
Dermatiseum thunbergii, while the lower flooding horizon was marked by
Acarospora schleicheri and species of the genus Heppia. In the tropics, similar
Coastal Lichens 7

with temperate latitudes, genera of marker species were recognized. McCarthy

(1999) described three species of Porina from aquatic and semiaquatic habitats in
this country and noted species of Staurothele as well as Verrucaria. In Papua New
Guinea (Aptroot 1998), four aquatic species of Verrucaria have been reported from
various heights (500–2100 m).
Recently, water lichens are becoming the objects of ecological and physiological
research. Coste et al. (2016) conducted a comparative study of photosynthetic
performance of epilithic lichens depending on different times of flooding. Obtained
results of this study showed slightly slower activation and higher sensitivity of PSII
to desiccation that may be important factors to explain the confinement of the most
freshwater-related species to habitats that provide sufficiently long hydration periods
(Coste et al. 2016).
Freshwater lichens were proposed as biological indicators of water level fluctu-
ations, water acidity, water quality (Gilbert and Giavarini 1997; Thüs et al. 2014),
and assessment of phosphate and nitrate pollution (Sonina 2014). However, their
potential as indicators needs further investigation in terms of sampling strategies to
obtain reproducible results and homogeneous interpretative scales (Thüs 2002;
Nascimbene and Nimis 2006).
Despite the fact that freshwater lichens, as previously reported, have not been
fully studied, a general trend is emerging – freshwater lichens correspond to rela-
tively few taxonomic groups (Nascimbene and Nimis 2006), and freshwater lichen
flora seems to be very homogeneous worldwide (Aptroot and Seaward 2003). This is
most likely determined by the fact that this group of organisms adapts to similar
conditions. The thalli of freshwater lichens are heavily physically affected by the
water stream (Thüs et al. 2014). In addition, frequent and prolonged periods of
complete flooding provide physiological stress hindering the diffusion of dissolved
gases such as oxygen and carbon dioxide inside the lichen thallus. Under such
conditions, thin crustose thalli are preferred, which facilitate gas exchange and
minimize physical exertion, as well as fruit bodies in the form of perithecia that
are immersed in the thallus (e.g., in Hydropunctaria) or surrounded by a protective
structure, such as involucrellum (most freshwater species with large perithecia).
Such morphological homogeneity in connection with the ecological limitation of
the environment is also confirmed by molecular studies performed for a limited
limitation (Thüs et al. 2015). Ecological limitations restrict morphological differen-
tiation and can partially explain the high degree of homoplastic characters of aquatic
lichens, as revealed by molecular studies, for example, for the Verrucariaceae family
(Gueidan et al. 2011).

3 Marine Lichens

Lichens of sea coasts are especially visible on rocks, where they can form a
continuous cover with 100% coverage (Fletcher 1980; Sonina 2014). Nevertheless,
this group of organisms is still poorly studied. Scattered references can be found in
8 A. V. Sonina and V. I. Androsova

works on marine fungi, where lichens are seen as a symbiosis with algae for the best
survival in saline conditions (Raghukumar 2017).
In contrast to freshwater lichens, the data on species diversity of marine lichens
have been largely restricted. On the coasts of 20 lagoons in England, the species
diversity of lichens on different types of rocky substrates has been studied, and
lichen flora is characterized by the presence of both specific coastal species and the
presence of upland species (Gilbert 2001).
For five localities on the territory of southwestern Greenland, 214 species of
lichens were found out, which differ in their distribution depending on the oceanic
climate (Hansen 2010). For the coasts of the White Sea and Barents Sea, 69 and 36
species were reported, respectively. Meanwhile this list included only epilithic
lichens that were found on boulders and rocks of the littoral and supralittoral (Sonina
2014).
Lichens from the Kandalaksha Bay of the White Sea are represented by 122
species from different substrates. Among these species, Verrucaria maura, V.
mucosa, Lichina confinis, and Caloplaca scopularis were listed as marine species
(Zhdanov and Dudoreva 2003). On the Baltic Sea coast in the southern part of the
Curonian Spit (the southern), 122 species of lichens from all types of substrates are
listed (Dolnik and Petrenko 2003). The authors have emphasized the fact that the
littoral and supralittoral zones differ significantly in species composition. For the
Pacific coast around Hong Kong, 27 lichen species are reported (Chu et al. 2000).
Researchers that studied sea coasts at different times reported clearly visible
lichen zones. The earliest works in which zones of sea coasts are noted belong to
the end of the nineteenth century (Grube and Blaha 2005). The study of the
principles of the lichen zonation on sea coasts and the ecological features of lichens
in different zones occur later. Some researchers examined lichen zones without
relating them to tidal rhythm (Lewis 1964), while others considered the importance
of tidal periodicity (Fletcher 1980; Hawksworth 1980; Chu et al. 2000; Zhdanov and
Dudoreva 2003; Gilbert 2003; Sonina 2014 etc.).
Fletcher carried out detailed studies on the rocky coast of Anglesey (Wales) and
identified four zones with subzones (Hawksworth 2000):

1. Sublittoral
2. Littoral (littoral subzone, Lichina pygmaea and Verrucaria mucosa, and littoral
border subzone – V. maura)
3. Supralittoral (three subzones: mesic, Caloplaca marina and Lichina confinis;
submesic, Xanthoria parietina; and xeric, Anaptychia runcinata and Ramalina
siliquosa)
4. Terrestrial (two subzones: halophilic lichens, Parmelia saxatilis, and halophobic,
P. omphalodes)

The height of the various zones mostly depends on the tidal rhythm and the type
of coast (rocks height and inclination angle of rocks), and the higher zones present
on the shores affected frequent severe storms (Hawksworth 1980). The similar
patterns of coastal lichens were reported in Hong Kong (Chu et al. 2000) for the
Coastal Lichens 9

Fig. 2 Tidal zone of the

White Sea coast

Pacific coast, on the White Sea coast in Russia (Sonina 2014). The study of the
Pacific coast of Hong Kong on coastal cliffs showed that coastal lichens (totally 27
species) are also distributed within four zones that differ in the lichen species
composition and the structure of lichen cover. The maximum projective cover of
lichens was observed in the 1st and 2nd zones while the greatest species diversity –
in the 3rd and 4th zones (Chu et al. 2000). The study also reported significant
differences in the species composition and distribution of lichens in the supralittoral
coastal ecosystems of the subtropical and temperate regions of the Northern Hemi-
sphere (Chu et al. 2000).
In studies of coastal lichens in the Barents Sea and White Sea (Figs. 2 and 3), two
ecological groups of lichens were recognized: halophytes and non-halophytes
(Sonina 2014). The author established lichen zone 4 on the upper intertidal and
supralittoral zones. Similar to other researchers, determination of lichen zones was
based on tidal rhythm, the presence of marker species, and structure of lichen
10 A. V. Sonina and V. I. Androsova

Fig. 3 Coast of the Barents

Sea

communities additionally. From zone 1 to 4 (from the waterline to land), the

structure of the lichen community changes from simple mono-species groups (e.g.,
Verrucaria ceuthocarpa or Hydropunctaria maura in zone 1) to complex multi-
species groups, which include lichens of different life forms (mainly non-halophytic
species in zone 4) (Sonina 2012).
According to results of ecological-coenotic analysis of coastal lichens of the
Kandalaksha Bay of the White Sea and based on characteristics of environmental
conditions, ten types of ecotopes on coastal rocks were recognized (Sonina et al.
2010). Stenotopic halophytic lichens occurred in four types of ecotopes from zones 1
and 2. These species inhabit in conditions of saltwater flooding. In the zones 3 and 4,
more favorable conditions are formed for the formation of epilithic lichen cover. Due
to larger diversity of habitats, six types of ecotopes were identified within these
zones, which were inhabited mainly by eurybiont species.
Coastal Lichens 11

Table 1 Сomparative analysis of coastal lichen species from different zones (measured by the
Jaccard index (KJ))
Lichen species
Zone The White Sea
1 zone Hydropunctaria maura, Verrucaria
ceuthocarpa, Wahlenbergiella mucosa
KJ ¼ 0.7
2 zone Acarospora amphibola, A. molibdina,
Caloplaca scopularis, Candelariella
vitellina, Ephebe lanata, Lecanora
helicopis, Physcia caesia, Pseudephebe
pubescens, Xanthoria candelaria,
Xanthoria elegans
KJ ¼ 0.3
3 zone Bellemerea alpina, B. cinereorufescens,
Candelariella vitellina, Catapyrenium
cinereum, Immersaria cupreoatra,
Lecanora dispersa, L. helicopis, L.
intricata, L. polytropa, L. lithophyla,
Lecidella anomaloides, Melanelia
hepatizon, M. stygia, Phaeophyscia
orbicularis, P. sciastra, Physcia caesia,
Rhizocarpon eupetraeum
KJ ¼ 0.2
4 zone Arctoparmelia incurva, Arctoparmelia
centrifuga, Diploshistes scruposus,
Lecanora intricata, L. polytropa,
Melanelia hepatizon, M. stygia,
Neofuscelia pulla, Parmelia saxatilis,
Physcia caesia, P. subalbinea, Porpidia
cinereoatr, P. crustulata, P. glaucophaea,
Rhizocarpon eupetraeum, R. geminatum,
R. geographicum, R. hochstetteri,
Umbilicaria deusta, U. hyperborea, U.
polyphilla, U. proboscidea, U. torrefacta,
Xanthoparmelia conspersa
KJ ¼ 0.3
The Barents Sea
Hydropunctaria maura, Verrucaria
ceuthocarpa
Caloplaca marina, Caloplaca saxicola,
Candelariella arctica, C. vitellina,
Lecania aipospila, Physcia caesia,
Xanthoria candelaria, Xanthoria elegans
Amandinea cacuminum, Lecania
aipospila, Lecanora intricata, L.
polytropa, Lecidea lapicida, L.
praenubila, Lobothallia melanaspis,
Melanelia hepatizon, Parmelia
omphalodes, Orphniospora moriopsis,
Physcia caesia, Rhizocarpon leptolepis,
Rinodina balanina, Rinodina milvina
Arctoparmelia centrifuga, Bellemerea
alpina, Clauzadea monticola, Lecanora
intricata, L. polytropa, Melanelia
hepatizon, Parmelia saxatilis, Parmelia
omphalodes, Protoparmelia nephaea,
Rhizocarpon geminatum, Rhizocarpon
geographicum, Rhizocarpon leptolepis,
Umbilicaria arctica, Umbilicaria
hyperborea, Umbilicaria proboscidea, U.
torrefacta

Sonina (2012, 2014), based on the analyzed literature as well as on the pairwise
comparison of partial lichen flora in the zones on the coasts of the Barents and White
Sea, suggested that the lichen cover of the coasts in the lichen zone 1 can be
attributed to the azonal type of lichen flora (Table 1).
According to the author (Sonina 2012, 2014), for the lichens of the lichen zone 1
on different continents, similar ecological and topographic conditions are formed
(periodic flooding with salt water and the presence of rocky substrate on the coast)
that are mostly related to tidal dynamics of the sea.
12 A. V. Sonina and V. I. Androsova

During the researches in the African continent on the Atlantic coast of the Namib
Desert the species lichen diversity was described, and the influence of stone surface
exposure on lichen distribution was shown (Wirth et al. 2007).
Despite the fact that lichens are widely represented on the coasts of all continents,
data on coastal lichens are scattered. The coastal lichens from temperate regions and
high latitudes of the Northern Hemisphere have been more extensively studied in
comparison with other territories.

4 Adaptations of Marine Lichens

The rocky sea coasts represent some of the most extreme habitats of living organ-
isms. Epilithic lichens are an essential component of the coastal ecosystems. They
occupy various ecological niches that are unsuitable for colonization by higher
plants. Lichens have to withstand the strong effect of abiotic factors, such as
fluctuations in solar radiation, waves and wind, daily drying and humidification
cycles, contrasting temperature, and salinity (Markovskaya et al. 2010; Sonina
2012). In the littoral zone, lichens need to adapt to the frequency of flooding with
salt water, while in the supralittoral zone, lichens withstand irrigation with sea
moisture and wave splash (Fletcher 1980). Only a limited number of studies have
attempted to factors influencing the distribution of lichens on rocky coasts. The main
factor determining the distribution and zonation of the phototrophic biota in the
coastal area is salinity (Delmail et al. 2013). Lichens growing on littoral and
supralittoral zones differ in their resistance to the duration of exposure to salt
water. Thus, the types of the littoral zone and supralittoral are clearly distinguished.
Species of the genus Wahlenbergiella occur in the littoral zone of the Northern
Hemisphere (Gueidan et al. 2009), and the species Verrucaria serpuloides in the
Southern Hemisphere grows constantly submerged in seawater (Lamb 1973), while
species of Hydropunctaria mainly occur in the supralittoral zone above the periodic
flooding zone (Gueidan et al. 2009; Orange 2012).
Some lichen species are not typical coastal species and widely distributed both on
the terrestrial area and on the coasts, such as Xanthoria parietina (Hawksworth
2000), and species of the genus Umbilicaria, which successfully colonize also stones
and rocks in forest communities, on the shores of lakes and rivers (Markovskaya et
al. 2010; Sonina 2014; Sonina and Tsunskaya 2016; Sonina et al. 2017). It remains
unclear what features determine the distribution in such different habitats. However,
the range of morphological and metabolic changes in these species is significant, and
further studies are required (Hawksworth 2000).
Studying the adaptations of halophyte lichens is fraught with difficulties because
lichens are complex organisms. It is important to understand the role that each biont
plays in lichen adaptation and the degree to which their responses to stress are
consistent (Delmail et al. 2013). The morphotype of lichen thallus, their ways of
reproduction, and their growth rate are reflected in their adaptive capabilities. The
secondary metabolites (lichen acids) are induced by extreme environmental condi-
tions and are involved in these processes. In studies of the lichen adaptation to the
Coastal Lichens 13

Fig. 4 Verrucaria
ceuthocarpa in the tidal zone
of the White Sea

conditions of sea coasts, morphological, structural, and functional adaptations are

distinguished.
Morphological adaptations. Morphological adaptive features were revealed for
the epilithic group of lichens of zones 1 and 2 on the coast of the Barents Sea (Sonina
2009): crustose densely cortical, scaly, and areolated life forms. Lichens of this
group are fused tightly to the substrate by hyphae of the medullary layer and resistant
to different events from the sea (from the zone 1 to 3). The peculiarity of the
reproductive strategy of coastal epilithic lichens is characterized by predominance
of sexual reproduction via ascospores. For littoral species, there is the presence of
perithecia – half-closed fruit bodies (species of the genus Verrucaria,
Wahlenbergiella) (Fig. 4). Small ascospores have more opportunities to settle and
hold on the surface of a stone substrate, in contrast to the larger particles of
vegetative propagation – soredia and isidia. Perithecia have a smaller contact area
between the water and spore-bearing part of the fruiting body.
Supralittoral species are more variable morphologically and anatomically. Mor-
phological and anatomical variability was revealed, and morphotypes were noted for
widespread supralittoral species on the coasts of the White and Barents Sea
Lecanora polytropa and Lecanora intricata (Sonina et al. 2011; Sonina 2014).
Such wide distribution of these species is connected with their ability to develop
various morphological, physiological, and biochemical adaptations, associated with
their responses to different climatic conditions. They have crustose thalli and are
14 A. V. Sonina and V. I. Androsova

White Sea Barents Sea

Lecanora intricata (Ach.) Ach.

Lecanora polytropa (Ehrh. ex Hoffm.) Rabenh

Fig. 5 Morphological adaptation of Lecanora intricata and Lecanora polytropa thalli in the
conditions of the White Sea and the Barents Sea costs

propagated by ascospores and/or conidiospores that can be fixed easily on the rocky
substrate on the coasts. However, the comparison of morphological features of these
species in the coasts of the Barents and White seas showed the existence of
morphotypes (Fig. 5). Lecanora intricata forms thinner thallus and unimmersed
apothecia on the White Sea coast.
Contrastingly, on the Barents Sea coast, the lichen’s thallus is thick, and the
apothecia are immersed in thalli. These peculiarities confirm the polymorphism of
this species. On the coast of the White Sea, Lecanora polytropa has an inconspic-
uous thallus with small dispersed areoles. In the condition of the Barents Sea, this
species has developed a thallus and submerged apothecia. On the White Sea coast,
this species spread more widely then along the Barents Sea coast. On the White Sea
coast, Lecanora polytropa occurs in the zones 2, 3, and 4 while on the Barents Sea
coast – zone 3 and 4, only. These species enhance its viability to the north. Analysis
of their anatomical structure showed that the total thickness of the thalli L. intricata
was significantly higher than thickness of the thalli L. polytropa. However, the ratio
of algal layer to the total thickness of the thallus was in all cases similar and was 1:3.
Coastal Lichens 15

Fig. 6 Umbilicaria
torrefacta on supralittoral of
the White Sea: habitation into
epilithic communities and
anatomy

In fact, morphological adaptations are mainly associated with the adaptability of

the mycobiont, which forms the thallus of the lichen reproductive structures and
provides direct contact with the environment.
Structural and functional adaptations. Structural and functional studies of
coastal lichen species were carried out on the White Sea coast (Sonina and
Tsunskaya 2016; Sonina et al. 2017). The objects of these studies were species of
the genus Umbilicaria and species Physcia caesia and P. dubia which belong to the
non-halophytic group and occurred in zones 3 and 4 (Figs. 6 and 7). Anatomical and
physiological characteristics (width of anatomical layers and the amount of photo-
synthetic pigments) of epilithic lichen species have been studied. The authors
reported two types of adaptations: structural due to the plasticity of the mycobiont
and functional due to the varying number of photosynthetic pigments of the photo-
biont. Ecological plasticity of the mycobiont which contacts the environment and
gives a niche to the photobiont has been revealed, based on the analysis of variance
16 A. V. Sonina and V. I. Androsova

Fig. 7 Physcia caesia on

supralittoral of the White Sea:
habitation into epilithic
communities and anatomy

in the study of anatomical structures of different lichen species and between different
samples of the same species. It was found that P. caesia adapts to the environmental
conditions through the variability of photosynthetic pigment levels which is con-
firmed by a strong variation of the chlorophylls a/b ratio and chlorophylls/caroten-
oids ratio with the stability of anatomical structures – functional adaptation. Lichen
adapts through the variability of anatomical layers and amounts of photosynthetic
pigments – structural and functional adaptation. For species U. torrefacta, the
structural adaptation to environmental conditions was recorded.
Despite the large body of literature on effects of salinity on plant physiology, little
is known concerning those factors driving the patterns of lichen functional charac-
teristics. In fact, high salt concentrations in the environment lead to a decrease in
hydric potential affecting water availability in poikilohydric organisms such as
Coastal Lichens 17

lichens (Hasegawa et al. 2000). Delmail et al. (2013) had gathered information about
ecology and mechanisms of adaptation to complex stressful environmental condi-
tions at cellular and biochemical levels in relation to symbiont diversity as well as
implications of phenotypic characteristics and genetics. In this detailed literature
review, authors have emphasized the halotolerance in lichens as a succession of
molecular and physiological events that lead to the regulation of hydric potential,
photosynthesis, osmolarity, and oxidative processes. Authors concluded that reac-
tions to salt stress at symbiotic stage are clearly different from those of isolated
partners. Each symbiont develops both specific arrays of metabolites for osmotic
adjustments and well adapted ionic mechanisms to limit nutrient imbalance, or it
activates certain antioxidant pathways involved in free radical and oxidative-com-
pound regulation (Delmail et al. 2013).
A number of studies documented an important role of photobiont, in the distri-
bution and adaptation to the coastal environment. The investigations by Ortiz-Á
lvarez et al. (2015) provide evidence that the photobiont, a strain of the
cyanobacterial photobiont Rivularia, played an important role in the distribution of
two species of the genus Lichina on the rocky shores of the Atlantic coast of Europe.
Two marine species Lichina pygmaea and L. confinis occur together on rocky
coasts but in different coastal zones. They are associated with the cyanobiont
Renularia. The authors found evidence of the high specificity of the cyanobiont
strains to both lichens species. Within each fungal species, selectivity varied across
the studied area, influenced by environmental conditions (both atmospheric and
marine), although patterns were highly correlated between both lichen taxa. Eco-
logical speciation due to the differential association of photobionts to each littoral
zone is suspected to have occurred in marine Lichina.
A pattern was found in relation to the photobiont species for littoral marine
lichens. They actually have no algae Trebouxia, Asterochloris, or Trentepohlia,
which are the most common among the photobionts of lichens. For marine species
from Verrucariaceae, a rather wide and unusual assortment of photobionts was
found (Thüs et al. 2011), including associations with Xanthophyceae (Parra and
Redon 1977), Phaeophyceae (Sanders et al. 2004), Trebouxiophyceae (Garrido-
Benavent et al. 2017), and Ulvophyceae (Thüs et al. 2011; Pérez-Ortega et al. 2018).
Confirmation of the important role of the photobiont in adapting to salinization
conditions of lichens was also registered in physiological experiments (Gasulla et al.
2019). In laboratory conditions, the results of osmoregulatory responses of algae
strains isolated from three lichen species were obtained: Hydropunctaria maura and
H. amphibia from the supralittoral and Wahlenbergiella striatula from the littoral.
The isolated strains belonged to the same species; Halofilum ramosum differed in the
ITS and RPL10A regions between the supralittoral H. maura and H. amphibia and
the littoral W. striatula species. The differences were in response strategies for high
salt concentrations: W. striatula accumulated glycerol, while H. maura and H.
amphibia synthesized sucrose. Analysis of the effectiveness of photosynthesis also
revealed differences: strains of algae from the thalli of W. striatula, living on the
littoral, showed lower photosynthetic activity under high irradiation (Gasulla et al.
2019).
18 A. V. Sonina and V. I. Androsova

Structural and functional adaptations of lichens represent the contribution of both

mycobiont and photobiont, which is responsible for the regulation of the synthesis of
various substances, including osmolytes, and are involved in the processes of lichen
colonization of the coastal zones.
Current interesting researches of coastal lichen species relate to the study of
bacteria associated with lichens. For terrestrial lichen species, such studies to
identify the bacterial group of lichen biota have long been carried out (Honegger
et al. 2013). However, there is still little known about bacteria of marine lichens. The
studies were performed on the littoral species Hydropunctaria maura (Bjelland et al.
2011). The bacterial communities of this lichen differed from the microbiota of
terrestrial lichens, by a high content of Actinobacteria, Bacteroidetes, Deinococcus,
and Chloroflexi. Notably, the role of these participants in the lichen association is still
not clear. Suzuki et al. (2015) reported that they produce secondary metabolites, and
some of these compounds have pronounced bioactive properties. A study of the
microbiota of other littoral species Lichina confinis and L. pygmaea (West et al.
2018) revealed differences between the bacterial communities associated with
marine and terrestrial lichens and even between marine lichens inhabiting different
coastal zones. The authors emphasized the importance of these studies for under-
standing the role of associated bacteria in the symbiosis of marine lichens and in
adaptation of this association to coastal environment.

5 Conclusion

The state of knowledge of epilithic lichens of sea coasts reflects the general state of
lichenological studies. Thus, studies of the species diversity of epilithic lichens have
been conducted, and general patterns of the formation of epilithic lichen cover on the
coasts have been revealed. It should be noted that these patterns are similar for
freshwater as well as for sea coasts, because the formation of a lichen cover and the
presence of lichen zones on the coast are associated with the effect of water bodies.
The morphological, anatomical, and functional adaptive features of lichen species to
coastal conditions have been established. As a result of recent research of the
physiological and molecular properties of lichen symbionts, an important role of
photobionts in the lichen adaptive strategies has been shown. Moreover developing
microbiological studies concerning lichens showed a specific role of microbiota in
their adaptation to certain coast habitats.
Nevertheless most of the fundamental issues in lichen distribution on the coasts
remain unresolved. Often, studies performed in these territories only induce new
questions that require reasonable answers. The complexity of their solution and the
excitement that they cause among researchers are limited to the symbiotic form of
existence of these fungi. It seems that the success of such studies will depend on a
combination of different research fields: floristic, environmental, physiological,
molecular, and genetic.
Coastal Lichens 19

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