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13Liverworts and mosses at the top of Elbrus (Caucasus) and bryophytes near limits of their altitudinal distribution
A. D. Potemkin1, G. Ya. Doroshina1, V. N. Mikhalenko2
1 Komarov Botanical Institute, Russian Academy of Sciences, St. Petersburg, Russia 2 Institute of Geography, Russian Academy of Sciences, Moscow, Russia
Corresponding author. A. D. Potemkin, [email protected]
Abstract. Two liverwort and two moss species were recorded from the outer side on the Eastern crater rim of Mt. Elbrus at 5590 m, on bare spot free of snow and ice about 10 x 20 m heated by geother-mal activity. They are Marsupella boeckii, which is recorded for the first time for Caucasus, M. cf. funckii recorded for the first time for Kabardino-Balkarian Republic, rare in forest belt in Caucasus Atrichum cf. angustatum and widespread Pohlia nutans. Their morphology, distribution and emergence at the top of Elbrus are discussed. Comparison of known data on the highest bryophyte records worldwide demonstrates their dissimilarity in different mountains. The bryophyte location at the top of Elbrus is highest in Russia and Europe. Despite it is lower than the known highest bryophyte records in the lower latitudes it is quite comparable with them due to more severe conditions at the top of Elbrus situated at 43°N.
Keywords: Marsupella boeckii, altitudinal distribution, bryophytes, Caucasus, geothermal activity, liverworts, mosses, Elbrus, Kabardino-Balkarian Republic, Russia.
Печеночники и мхи на вершине Эльбруса (Кавказ) и мохообразные близ границ их высотного распространения
А. Д. Потемкин1, Г. Я. Дорошина1, В. Н. Михаленко2
1 Ботанический институт им. В. Л. Комарова РАН, Санкт-Петербург, Россия 2 Институт географии РАН, Москва, Россия Автор для переписки. А. Д. Потемкин, [email protected]
Резюме. Два вида печеночников и два вида мхов были собраны на наружном крае кратера восточной вершины Эльбруса, на высоте 5590 м, на лишенном снега и льда участке, площадью примерно 10 x 20 м, обогреваемом за счет геотермальной активности. В этом местообитании обнаружены Marsupella boeckii, выявленная впервые для Кавказа, M. cf. funckii, приводимая впервые для Кабардино-Балкарской Республики, редкий в лесном поясе на Кавказе Atrichum cf. angustatum и широко распространенный вид Pohlia nutans. Обсуждены морфология, распространение и появление на вершине Эльбруса этих видов. Обобщены и проанализированы в сравнительном плане данные по самым высокогорным находкам мохообразных в мире. Показано, что видовой состав мохообразных, выявленных на самых высоких горах, различен. Местонахождение мохообразных на вершине Эльбруса самое высокое в России и Европе. Несмотря на то, что оно расположено несколько ниже, чем самые высокие местонахождения мохообразных в значительно более низких широтах, оно вполне сопоставимо с ними в связи с более суровыми условиями на вершине Эльбруса, расположенного на 43° с. ш.
Ключевые слова: Marsupella boeckii, высотное распространение, геотермальная активность, мохообразные, мхи, печеночники, Кабардино-Балкария, Кавказ, Россия, Эльбрус.
Introduction
Bryophytes are the most widespread land plants. They occur from high polar latitudes to highest elevations. Their taxonomic composition, abundance and participation in plant cover formation are quite specific in high mountain areas, high Arctic and Antarctic regions (Halloy, 1991; Seppelt, Ochyra, 2008; Afonina, 2015; Potemkin, Matveyeva, 2015).
Occurrence of bryophytes near the limits of their distribution is caused by the presence of suitable habitats which are free of snow and ice for several months per year and have accessible water first of all. There are two different types of such habitats near the limits of bryophyte distribution: cold habitats and those heated by volcanic activity. Both of them occur in high latitudes and high mountains. They are still insufficiently studied because of inaccessibility.
The goal of this communication is to report the highest bryophyte records in Europe and Russia, to make their comparison with the previously known highest bryophyte records in the world, and to discuss the known data on bryophytes near the limits of their distribution.
Material and methods
In 2014, bryophytes were collected by glaciologists at the outer side of the crater rim of the Mt. Elbrus at 5590 m of altitude on a bare spot free of snow and ice of about 10 x 20 m heated by geothermal activity. The collecting place is strongly isolated from the vegetation beneath by extensive permanent snow and ice fields about 1.5-2 km high. According to three-year (2013-2016) air temperature records at the eastern summit of the Mt. Elbrus (5630 m), the current temperature data have never reached positive values. Minimum air temperature (-39.8 °C) was measured on December 11, 2013. Maximum one (-6.3°C) was registered on August 15, 2014. At the same time, due to the geothermal heat flux, the temperature at the sampling site surface is always positive and varies from 17.0 °C to 26.5 °C. At a 40-cm depth below surface, the range of temperature variability is even smaller and varies from 25.6 °C to 27.6 °C (Mikhalenko et al, in prep.). This provides the necessary amount of heat and liquid moisture for the year-round existence of these species. Detailed characteristic of the sampling site is given in a separate paper (Mikhalenko et al., in prep.).
The moss cover is fragmentary at the collecting site (Plate I: 2, 3). According to available photographs and collected specimens Atrichum cf. angustatum and Marsupella boeckii are most abundant in the site and form extensive continuous cover of the substrate, whereas Marsupella cf. funckii and Pohlia nutans associate with them as groups of plants. Bryophytes were found in seven collecting bags. They were studied with traditional morphological methods and modern taxonomic literature. Micrographs were taken with an Olympus Stylus Tough TG-3 digital camera using its stacking mode. Methods used for description of collecting sites are described in a separate paper (Mikhalenko et al., in prep.).
Plate I. 1 — Eastern crater of Elbrus, 2, 3 —collecting place of bryophytes at the outer side of the Eastern crater rim of Elbrus at 5590 m, 4 — Aulacomnium cf. angustatum in collecting place.
Photo by V. N. Mikhalenko.
Plate II. Aulacomnium cf. angustatum (1-5), Pohlia nutans (6-10), Marsupella cf. funckii (11-14), M. boeckii (15-20) from the top Elbrus. 1, 8-11, 15, 16 — shoots and shoot sectors; 2, 12, 13, 17, 19 — leaves; 3, 4 — leaf cross sections; 5, 6, 7 — upper parts of leaves; 14, 20 — stem cross sections; 18 — leaf cells. Scale bars: 1, 8, 10 — 1 mm, 2 — 85 ^m, 3 — 18 ^m, 4, 5, 7 — 75 ^m, 9 — 200 ^m, 11, 12 — 100 ^m, 13 — 60 ^m, 14, 20 — 250 ^m, 15, 16 — 120 ^m. All from 4 VIII 2014, Mikhalenko, LE. Photo by A. D. Potemkin.
Results
Liverworts
Marsupella boeckii (Austin) Lindb. ex Kaal. (Plate II: 15-20) — Outer side of the East-em crater rim of the Mt. Elbrus, on bare volcanic soil, 43°20'58.7"N, 42°27'14.5"E, 4 VIII 2014, Mikhalenko, LE, 5590 m alt. It often forms extensive continuous cover, sometimes with admixture of Atrichum cf. angustatum and Pohlia nutans. Sterile. Studied plants have rather typical morphology, represented by mod. parviretis-colorata trans. ad viridis and distinguished by sporadic purplish rhizoids.
Infrequent Holarctic species known from Europe, Asia and North America (Damsholt, 2002). In Russia it is known from Kola Peninsula, North Urals, South Siberia and Far East (Potemkin, Sofronova, 2009). First record for Caucasus and Kabardino-Balkarian Republic. In Europe, this liverwort occurs at lower altitudes, in Eastern Alps it was recorded at 1900-2400 m (Müller, 1951-1956), in Bhutan at 3640 m (Grolle, Long, 1990). In Russia, it was recorded from North Taiga to Arctic Tundra, in mountain taiga and tundra communities (Potemkin, Sofronova, 2009).
Marsupella cf. funckii (F. Weber et D. Mohr) Dumort. (Plate II: 11-14) — Outer side of the Eastern crater rim of the Mt. Elbrus, on bare volcanic soil, 43°20'58.7"N, 42°27'14.5"E, 4 VIII 2014, Mikhalenko, LE, 5590 m alt., associated with Marsupella boeckii, Pohlia nutans and Atrichum cf. angustatum.
M. funckii has chiefly European distribution with isolated locations in Turkey, eastern North America, and Japan (Damsholt, 2002). In Russia besides Caucasus it was recorded from the Kaliningrad Region, the Republic of Karelia, South Siberia and Far East (Potemkin, Sofronova, 2009; Potemkin et al., submitted). It is known in Caucasus up to 3000 m alt. from Teberdinsky Reserve in the Karachayevo-Circassian Republic (Ignatova et al, 2008) and Caucasian State Nature Reserve in the Republic of Adygea (Konstantinova et al, 2009). Studied plants are poorly developed but distinct from M. boeckii in broader patent leaves with often deeper sinuses which provide principally different habit of plants, which corresponds well to impoverished plants illustrated by Müller (1951-1958: Fig. 260).
Mosses
Atrichum cf. angustatum (Brid.) Bruch, Schimp. et W. Gümbel (Plates I: 3-4; II: 1-5) — Outer side of the Eastern crater rim of the Mt. Elbrus, on bare volcanic soil, 43°20'58.7"N, 42°27'14.5"E, 4 VIII 2014 Mikhalenko, LE, 5590 m alt., among Marsupella boeckii, partly with Pohlia nutans. Strongly impoverished sterile plants abundantly developed in the spot.
Known from Europe, Macaronesia, Turkey, Iran, eastern and central North America and China (Atrichum angustatum, EOL site). In Caucasus, it has been previously recorded in a
forest belt only (Doroshina, unpubl.) In Europe, it was recorded from 900 to 1700 m alt. (Lim-pricht, 1895) and probably higher, in China from 1500-2500 m alt. (Wu, Wang, 2005).
Pohlia nutans (Hedw.) Lindb. (Plate II: 6-10) — Outer side of the Eastern crater rim of the Mt. Elbrus, on bare volcanic soil, 43°20'58.7"N, 42°27'14.5"E, 4 VIII 2014 Mikha-lenko, LE, 5590 m alt., as an admixture to Marsupella boeckii, together with Atrichum cf. angustatum. Sterile plants with typical morphology.
Widespread species known from Europe, Asia, North America, South Africa, Australia, Antarctica (Zhang et al., 2007). In Caucasus it was found up to alpine belt (Doroshina, unpubl.). In Europe, it was recorded up to 2530 m alt. (Limpricht, 1895) and probably higher. In China, it was recorded from 3000-4000 m alt. (Zhang et al., 2007).
Discussion
Emergence of bryophytes at the top of Elbrus. Discoveries of bryophyte species on high mountains are often difficult to explain. It may result from transfer of diaspores by airflow, alpinists and in some cases from migrating birds. Highest bryophyte populations may be recruited from lower altitudinal belts or, in some cases, from different unidentified sources (see: Halloy 1991).
The emergence of bryophytes at the top of Elbrus is not clear. The revealed taxonomic composition includes a widespread Pohlia nutans, known from a few sites in Caucasus Marsupella cf. funckii and Atrichum cf. angustatum (the latter from forest belt only) and Marsupella boeckii, which has not been previously recorded from Caucasus. The revealed bryophyte species composition at the top of Elbrus is quite specific for Caucasus. It is noteworthy that all the species in question have small anemochorous spores, 7-18 to 16-22 ^m in diam., that makes possible their transfer to the top of Elbrus by strong winds. As it was mentioned by Zanten and Pocs (1981: 504) «particles with a radius of up to 10 or 20 microns can be found at the altitude of up to 5 or 6 km, but it is unlikely that they reach greater heights». It seems possible that three or all four species were transferred to the top of Elbrus by strong winds. Marsupella cf. funckii and Pohlia nutans could be brought to the top of Elbrus by alpinists, too. It should be mentioned that Ceratodon pur-pureus (Hedw.) Brid., which is very common along alpinist trails up to snow level on Elbrus, has not been recorded in collections from the top of Elbrus.
High altitude bryophyte distribution. Analysis of annotated lists of high elevation territories and taxonomic accounts, which include data on elevation range of the studied taxa, usually provides rather limited information on the species altitudinal distribution. This makes clear that our data on elevations, where bryophytes occur, are much limited due to frequent inaccessibility of such locations for bryophyte collectors. The bryophyte distribution and species composition in high mountains are caused by latitudinal position
of mountains, structure and composition of bedrocks, continentality of climate of the studied territory, microclimate features and temperature of inhabited substrates, air humidity and/or accessibility of water and their constancy in the bryophyte habitats.
In tropical areas bryophyte species from different taxonomic groups occur and often common in the areas as high as 5000-5100 (5200) m (Grolle, 1966; J. Shevock, pers. comm. for Sichuan, 11 February 2016). This points at quite possible much higher distribution of bryophytes in those areas. Discoveries of mosses at much higher altitudes (Table)
confirm this assumption.
Table
Highest bryophyte locations at the world
Bryophyte taxon Highest record Mount Coordinates Location Substrate temperature Source of information
Mosses
Aongstroemia 6035 m Everest, ~ 28°06'N Asia, unknown Dixon,
julacea (Hook.) Mitt. (19800 ft) Rong- buk Valley 86°51'E Nepal 1925
Atrichum cf. 5590 m Elbrus 43°20'58.7'N, Caucasus, 17.0 - 26.5°C Present
angustatum 42°27'14.5"E (43°20.979N, 42°27.241'E) Russia at the surface 25.6 - 27.6°C at 40-cm depth report
Bryum pallens 5760 m Hindu 35°N, 71°E Middle unknown Dixon,
Sw. (18900 ft) Kush Asia, Afghanistan 1924
Bryum sp. 6060 m Sacompa 24°25'S, 68°15W South America, Argentina 31°C at 10-cm depth Halloy, 1991
Globulinella 6030 m Sacompa 24°25'S, South 31°C at 10-cm Schiavone,
halloyi Schia- 68°15W America, depth Suarez,
vone et Suárez Argentina 2009
Grimmia som- 5029- Everest, ~28°05'60" N Asia, near unknown Dixon,
ervellii Dixon 5334 m (1650017500 ft) near Lamna La Pass 86°39E ChinaNepal border 1925
Gymnostomum 6480 m Everest unknown Asia unknown Sym-
aeruginosum Sm. biosis, sites
Hygroamblys- 5761 m, Hindu 35°N, 71°E Middle unknown Dixon,
tegium filicinum (Hedw.) Loeske cf. var. supra-alpinum (Mo-lendo) Glow. (18900 ft) Kush, Asia, Afghanistan 1924
Orthotrichum 5791 m, Chim- 1°28'9"S, South unknown Dixon,
wagneri Dixon (19000 ft) borazo 78° 49' 3" W America, Ecuador 1924
Table (continued)
Bryophyte taxon Highest record Mount Coordinates Location Substrate temperature Source of information
Pohlia sp. 6060 m Sacompa 24025'S 68015'W South America, Argentina 31°C at 10-cm depth Halloy, 1991
Pohlia nutans 5590 m Eastern peak of Elbrus 43020'58.7"N, 42027'14.5"E (43020.979'N, 42027.241'E) Caucasus, Russia 17.0 - 26.5°C at the surface 25.6 - 27.6°C at 40-cm depth Present report
Liverworts
Marchantia berteroana Lehm. et Lindenb. 6060 m Sacompa 24025'S, 68015'W South America, Argentina 310C at -10 cm Halloy, 1991
Marsupella boeckii 5590 m Eastern peak of Elbrus 43020'58.7"N, 42027'14.5'E (43020.979'N, 42027.241 E) Caucasus, Russia 17.0 - 26.50C at the surface 25.6 - 27.60C at 40-cm depth Present report
Marsupella cf. funckii 5590 m Eastern peak of Elbrus 43020'58.7"N, 42027'14.5'E (43020.979'N, 42027.241 E) Caucasus, Russia 17.0 - 26.50C at the surface 25.6 - 27.60C at 40-cm depth Present report
In most high mountain areas, bryophyte uphill distribution is usually limited by the level of permanent snow. In high altitudes, their findings are possible in snow-free areas, which may occur in some hidden from snowfalls places like it was found on the Mt. Everest at 6480 m alt. (Symbiosis on Mt. Everest, sites) and in high dormant volcanoes due to their geothermal heat flux causing snow melting close to their summits. The principal factors affecting bryophytes in such extreme conditions are short snow-free period, soil temperature and humidity, UV-radiation, and, according to Halloy (1991), CO2 concentration in air at ground level. These factors hamper sexual reproduction of bryophytes and form a barrier for their distribution in highest mountains. However, asexual reproduction of bryophytes usually takes place under such unfavorable conditions and results in survival of their populations in such habitats.
Comparison of bryophyte species composition known from the highest mountains has shown its dissimilarity in different mountains. The following species were recorded from highest altitudes of different mountains: Aongstroemia julacea, Grimmia somervellii, Gym-nostomum aeruginosum from Everest, Himalaya, Orthotrichum wagneri from Chimboradzo, South America, Bryum pallens, Hygroamblystegium filicinum from Hindu Kush, Middle Asia, Globulinella halloyi, Bryum sp., Pohlia sp., Marchantia berteroana from Sacompa, Andes, and Atrichum cf. angustatum , Pohlia nutans, Marsupella boeckii, M. cf. funckii from Elbrus, Caucasus (Table). Acrocarpous mosses most often occur in highest altitudes and only three liverwort species, two leafy species of Marsupella Dumort. and complex thallose
Marchantia L., were found in highest mountains at 5590 and 6030 m, all in places with warm substrates due to geothermal heating. According to photograph of highest bryophyte location of Gymnostomum aeruginosum from Everest, at 6480 m, it is hidden in crevice among snow and should be cold (Symbiosis on Mt. Everest, sites). Predominance of acrocarpous mosses in the list in the table points at their best adaptation for such unfavorable conditions in high altitudes in comparison with liverworts.
Conclusions
Known bryophyte records in highest elevations of different mountains are dissimilar and recruited from local and remote floras.
The bryophyte records from the top of Elbrus are highest in Russia and Europe. Despite they are lower than the known highest locations in lower latitudes, they are quite comparable with them due to more severe conditions at the top of Elbrus.
Isolated bryophyte populations at the top of Elbrus represent a remarkable model for monitoring, which reflects climate changes, possibilities for their introduction from the lower altitu-dinal belts, morphological and genetic transformations of bryophytes resulted from such a high position above the sea level. It is recommended to demarcate the recorded spot and to make its annual photodocumention (preferably, macrophotographs) together with collecting small specimens of morphologically different plants for their identification by specialists.
Acknowledgments
James Shevock is thanked for personal communication on his observations on bryophytes at high altitudes in Sichuan. John Brinda, Caspar Chater, Juan Larrain, Genevieve E. Tocci are acknowledged for sharing valuable internet resources; help of Tamas Pocs with the literature is appreciated. We are grateful anonymous reviewers and Yuri Okolod-kov for reading the manuscript, valuable advices and correcting the English. The study of A. D. Potemkin and G. Ya. Doroshina was carried out within the framework of the institutional research project 01201255616, the study of V. N. Mikhalenko was supported by Russian Science Foundation Grant 17-17-01270.
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Halloy S. 1991. Islands of life at 6000 m altitude: the environment of the highest autotrophic communities on Earth (Socompa Volcano, Andes). Arctic Alpine Res. 23(3): 247-262.
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