The distribution of aquatic mosses among 73 lakes in the Soya Coast region, East Antarctica, was surveyed. Two species of mosses, Bryum pseudotriquetrum and Leptobryum sp. were found at the bottom of lakes. B. pseudotriquetrum was found in 38 lakes (52.1%), mainly in freshwater lakes throughout the study area. Leptobryum sp. was found in 26 lakes (35.6%) in a rather restricted area, and mainly in relatively saline lakes

Hiroshi Kanda

Koji Seto

Sakae  Kudoh

Satoshi Imura

Shuji Ohtani

Tadashi Bando

National Institute of Polar Research Repository

Distribution of aquatic mosses in the Soˆya Coast region,East AntarcticaSatoshi Imura+, Tadashi Bando,, Koji Seto-, Shuji Ohtani.,Sakae Kudoh+ and Hiroshi Kanda++National Institute of Polar Research, Kaga, +-chome, Itabashi-ku, Tokyo +1-2/+/,Department of Biology, Kyoto University of Education, Fukakusa, Fushimi-ku,Kyoto 0+,2/,,-Research Center for Coastal Lagoon Environments, Shimane University,+*0*, Nishi-kawatsu, Matsue 03*2/*..Department of Education, Shimane University, +*0*, Nishi-kawatsu, Matsue 03*2/*.Abstract: The distribution of aquatic mosses among 1- lakes in the Soˆya Coastregion, East Antarctica, was surveyed. Two species of mosses, Bryum pseudo-triquetrum and Leptobryum sp. were found at the bottom of lakes. B. pseudotriquetrumwas found in -2 lakes (/,.+), mainly in freshwater lakes throughout the study area.Leptobryum sp. was found in ,0 lakes (-/.0) in a rather restricted area, and mainlyin relatively saline lakes.key words: Antarctica, lake, aquatic moss, distribution, salinityIntroductionBryophytes occur in a diverse range of terrestrial and also aquatic habitats.Ignatov and Kurbatova (+33*) reviewed the occurrence of aquatic bryophytes, especial-ly in deep lakes in the world, and listed about 2/ taxa of mosses and liverworts. Theseaquatic bryophytes, which grow on bottom substrata in lakes and streams, and perma-nently in submerged condition, have often been found in high altitude (e.g., Light andSmith, +310) or high latitude regions (e.g., Longton, +322), where the environmentalconditions are very severe for macrophytes and aquatic seed plants.In the Antarctic region, +/ taxa of mosses are known as aquatic plants at thebottom of lakes (Table +). In the Maritime Antarctic, 0 taxa of mosses have beenreported in a large number of lakes on Signy Island (Light and Heywood, +31-; Priddleand Dartmall, +312; Priddle, +313), and / taxa of mosses are known from lakes onAlexander Island (Light and Heywood, +31/). Even in some ice free areas in EastAntarctica, the most severe terrestrial habitat on Earth, / taxa of aquatic mosses havebeen reported (e.g., Savich-Lyubitsukaya and Smirnova, +3/3; Kasper et al., +32,;Seppelt, +32-; Kanda and Iwatsuki, +323). Light and Heywood (+31/) emphasizedthe luxuriance of aquatic moss communities in the lakes, contrasting strongly with thepaucity of terrestrial vegetation in East Antarctica.In the Soˆya Coast region, four taxa of aquatic mosses have been reported. TheyE-mail: imura@nipr.ac.jp+Polar Biosci., +0, ++*, ,**- ,**- National Institute of Polar Researchare Bryum pseudotriquetrum (e.g., Kanda and Ohtani, +33+), Bryum sp. (Kanda andMochida, +33,), Henediella heimii (Kanda and Ohtani, +33+ as Pottia heimii) andLeptobryum sp. (e.g., Imura et al., +333; Kanda and Iwatsuki, +323 as Dicranella sp.).Most of these works on aquatic mosses are reports on their occurrences in particularlakes, or taxonomical treatments. Only Kanda and Mochida (+33,) discussed thedistribution of aquatic mosses in a study at Skarvsnes, one of the ice free areas in thisregion.Imura et al. (+333) described a curious aquatic moss vegetation, described as mosspillars, in some lakes in the vicinity of Syowa Station. Subsequently, a more detailedsurvey of the aquatic mosses in many lakes in the Soˆya Coast region, East Antarctica,was carried out. In this report, we compile the results of our surveys to clarify thedistribution of aquatic mosses in this region, and discuss the limiting factors governingtheir distribution.MethodsThe Soˆya Coast is located on the east coast of Lu¨tzow-Holm Bay, Dronning MaudLand, East Antarctica (Fig. +). Syowa Station is situated on East Ongul Island, at thenorth end of this region. Along the +** km long coast from West Ongul Island (ca. 03S)at the entrance of Lu¨tzow-Holm Bay, to Strandnibba (ca. 1*S) in the innermost partof the bay, several ice-free areas and islands are interspersed. Seven ice-free areas wereTable +. List of the aquatic mosses in Antarctic lakes.Aquatic mosses Signy Island Alexander Island East AntarcticaAmbrystegium sp.Calliergon sarmentosumCampylium sp.Drepanocladus sp.Drepanocladus cf. aduncusPohlia nutansBryum sp.Campylium polygamumDicranella sp.Distichium capillaceumBryum pseudotriquetrumBryum sp.Leptobryum sp.Henediella heimiiPlagiothecium simonowii++, ,, -+++, ,+..... /+,+++++0+*+1+: Light and Heywood (+31-), ,: Priddle (+313), -: Priddle and Dartmall (+312), .: Lightand Heywood (+31/), /: Savich-Lyubitskaya and Smirnova (+3/3), 0: Savich-Lyubitskayaand Smirnova (+30*), 1: Kasper et al. (+32,), 2: Seppelt (+32-), 3: Kanda and Iwatsuki(+323), +*: Kanda and Ohtani (+33+), ++: Kanda and Mochida (+33,), +,: Imura et al.(+333), +-: Nakanishi (+311), +.: Ochi (+313), +/: Imura et al. (+33,), +0: Tewari and Pant(+330), +1: Savich-Lyubitskaya and Smirnova (+30.).S. Imura et al.2surveyed: West Ongul Island, the Langhovde area (including Hamnenabben), Breiva˚g-nipa, the Skarvsnes area (including Byva˚ga˚sane), Skallen, Rundva˚gshetta andStrandnibba.On the ice free area in this region, there are many lakes of various sizes, depths andwater qualities. Lakes over +m in depth were selected in this study. In total, 1- lakeswere surveyed from +33. to ,**+, during the -0th to .,nd Japanese Antarctic ResearchExpeditions. The name, geographical characteristics, and electric conductivity of lakessurveyed in this study are listed in Table ,. A directory and maps of each lake areshown on the web page (http: //www.isc.nipr.ac.jp/penguin/Terrestrial/regal/DataBase/index.htm).The benthic vegetation of lakes was collected in several ways. When the lakesFig. +. Map of the study area.Distribution of aquatic mosses 3Table,.Geologicalandchemicalcharacteristicsofsurveyedlakesanddistributionofaquaticmosses.S. Imura et al.4Distribution of aquatic mosses 5were ice-free, we used a rubber boat and an Ekman grab sampler or a rake with longhandle to collect the vegetation. When the lakes were frozen, we made holes in the ice,and used the rake through the ice hole. In winter, we collected the mass of benthicvegetation exposed on the ice-surface of lakes. This vegetation was considered to havebeen growing at the bottom of lakes before being moved upward through the ice cover(Wilson, +30/). All samples were frozen at ,*C and carried back to the laboratory,and observed by light microscope.As the most basic measure of water quality, electric conductivity (EC) of lakewater was measured in situ at some lakes. From each vertical proﬁle of EC values, thedeepest but in maximum -m in depth value was used, because the aquatic mosses wereknown to be most vigorous at -m in depth in this region (Imura et al., +333).Results and discussionTable , shows the distribution of aquatic mosses in 1- lakes surveyed in the SoˆyaCoast region. The aquatic vegetations of mosses were found in .+ lakes (/0.,). Inthe Soˆya Coast region, aquatic mosses are thought to be common components of the lakebottom vegetation. These vegetations were found at the bottom of lakes from ,.* to+../m in depth, and were most vigorous from - to /m. In shallow areas, less than ,m in depth, moss vegetations were not found, probably due to ice action, there were onlyfragments of algal mats. In the surveyed lakes, the maximum thickness of surface icein winter was less than ,m. Bodin and Nauwerck (+302) noted that ice movement insummer is the main reason to explain the absence of aquatic moss colonies in the shallowwater zone down to about ,m depth. The deepest record of aquatic mosses in EastAntarctica is -,.-m, where Plagiothecium georgico-antarcticum was found in LakeGlubokoye, Schirmachar Oasis, Dronning Maud Land (Savich-Lyubitskaya andSmirnova, +30.). The world’s deepest record is +/-m, where mosses and liverwortshave been found in Lake Tahoe, California-Nevada (Frantz and Cordone, +301). Inthis study, the maximum depth for aquatic mosses was just +../m, but there were onlya few lakes deeper than this depth, and most of them were hyper saline lakes.In this study, we found two species of mosses, Bryum pseudotriquetrum (Hedw.)Gaertn., Meyer et Scherb. and Leptobryum sp. at the bottom of many lakes in the SoˆyaCoast region. Bryum sp. andHenediella heimii were not found in the lakes surveyed inthis study. Aquatic Bryum sp. was reported from two lakes in Skarvsnes by Kanda andMochida (+33,), but it has not been reported since then. An aquatic form ofHenediella heimii was reported from lake Yukidori Ike in Langhovde by Kanda andOhtani (+33+) as Pottia heimii. In this study, ﬂoating submerged shoots of H. heimiiwere sometimes found in the lakes, but not on the bottom substrata. It is suggested thatﬂoating shoots of the species are temporarily submerged from the population growing onthe soil around the lake. We have therefore excluded Bryum sp. and H. heimii fromthe list of aquatic mosses in the Soˆya Coast region.Bryum pseudotriquetrum has been reported in lakes in several regions in Antarctica,such as Alexander Island (Light and Heywood, +31/ as B. algens), the Dry Valleyregion (Kasper et al., +32, as B. algens), Vestfold Hills (Seppelt, +32-) and BungerHills (Savich-Lyubitskaya and Smirnova, +3/3 as B. korotkevicziae). The morpholog-S. Imura et al.6ical characteristics of aquatic B. pseudotriquetrum have been well described and il-lustrated by Seppelt (+32-), Kanda and Iwatsuki (+323) and Kanda and Ohtani (+33+).This species is the commonest moss around ephemeral streams in Antarctica, and alsovery common and widespread, mainly in wet conditions around the world.Irregular mat-shaped vegetations of B. pseudotriquetrum were found in -2 of the 1-lakes (/,.+). This species was not found on West Ongul Island, but was found in allof the other areas from Langhovde (ca. 03+*S) near the entrance of Lu¨tzow-HolmBay to Strandnibba (ca. 1*S) in the innermost part of the bay. In the Skarvsnes area,it was found in ,2 of .. lakes surveyed (0-.0). B. pseudotriquetrum is thought to bea quite common and widespread species at the bottom of lakes in this region.Leptobryum sp. at the bottom of lakes in this region was ﬁrst reported by Nakanishi(+311) as Bryum korotkevicziae Sav. et Smirn. or its variety hollerbachii Sav. et Smirn.Ochi (+313) treated it as Bryum sp., and Kanda and Iwatsuki (+323) considered it to beDicranella sp. Imura and Kanda (+32/) and Imura et al. (+33,) investigated themorphology of tubers (vegetative diaspores), and identiﬁed it as Leptobryum pyriforme(Hedw.) Wils. Imura et al. (+333) argued that the taxonomic position of this specieswas still uncertain, and treated it merely as Leptobryum sp. Recently, Arts (,**+)reviewed the genus Leptobryum around the world, and considered the aquaticLeptobryum in Antarctica to be L. wilsonii (Mitt.) Broth. based on the illustration ofaquatic plants by Kanda and Iwatsuki (+323). He described the sexuality of L.wilsoniias dioicous, while Imura et al. (+33,) reported synoicous inﬂorescences on shoots fromcultured tubers of the aquatic Leptobryum. Further morphological and moleculargenetic studies are needed to clarify the taxonomic position and origin of aquaticLeptobryum sp. in Antarctica.Leptobryum sp. was found in ,0 of 1- lakes surveyed (-/.0). It was foundmainly in the Skarvsnes area (,. of .., /../) and only one lake in the Langhovde area(the neighbor to the north of Skarvsnes) and Skallen (the neighbor to the south ofSkarvsnes). Leptobryum sp. is relatively rare at the bottom of lakes in this region, andthe distribution area is restricted mainly to Skarvsnes. Imura et al. (+333) reportedcurious pillar-like vegetation of this species at the bottom of some lakes in Skarvsnes.Also in this study, moss pillars were found in 2 lakes only in Skarvsnes (Table ,). InLanghovde and Skallen, Leptobryum sp. formed rough irregular mat-shaped vegetation.Table - shows the occurrence of two aquatic mosses among /* lakes for whichelectric conductivity (EC) data were obtained. B. pseudotriquetrum was found in lakeswith quite low (+.*mS/m, lake Maruwan O«ike) to relatively high (,+*.* mS/m, lakeHyotan Ike) EC values; it was most vigorous in low salinity lakes. On the other hand,aquatic Leptobryum sp. was not found in lakes with very low EC values, but fromrelatively low (+/.*mS/m, lake Dojyou Ike) to high (-3*.*mS/m, lake A-1 Ike) values.This highest EC value of lake A-1 Ike is close to one-tenth of the value in seawater.It is noteworthy that Leptobryum sp. has never been found in terrestrial habitats inEast Antarctica. Except in the Syowa Station area, this species has been reported onlyin some lakes around the Indian Maitri station (1*-1S, *2,,E) at SchirmacharOasis, over +*** km west of Syowa Station (Tewari and Pant, +330). Smith (+32.)reported the occurrence of this genus from Deception Island (0-**S, 0*.*E) in theSouth Shetland Islands, Maritime Antarctic, and this is the only report of this speciesDistribution of aquatic mosses 7Table -. Electric conductivity of lakes and distribution of aquatic mosses.LakesElectricconductivity(mS/m)Aquatic mossesBryum pseudotriquetrum Leptobryum sp.Higasi-Hamna IkeMaruwan O«ikeGokuh IkeMaruwan-minami IkeMaruwan-kita IkeYukidori IkeSt-, IkeTenno-kama IkeKami-tenpyoˆ IkeHyouga IkeNaka-tenpyoˆ IkeSt-+ IkeKo IkeX IkeDojyou IkeKoke NumaKuwai IkeKami-kama IkeHigasi-yukidori IkeA-. IkeSkallen O«ikeRokuban IkeO«-ikeNamazu IkeMisumi Ike*40+4*-4/-40.4*.4*.4+.4..40/4*14,141242+.4.+/4*+/40+04*+14++143+24*+24.+242,-41,.4*,34/B-, IkeB-+ IkeMedaka IkeOyako IkeTanago IkeB-- IkeA-/ IkeShimo-tenpyoˆ IkeNamagi IkeB-. IkeNaga-IkeA-- IkeA-0 IkeA-2 IkeHyotan IkeA-, IkeA+* IkeMagobati IkeAkebi IkeA-1 IkeKobati IkeNurume IkeZakuro IkeOyayubi IkeSuribati Ike-*4*..4*/-4//.4./34/0,4*1/42104/3341+*,4*++04,+,*41+1.4*,**4*,+*4*,.*4-,3*4*,3*4+-1.4/-3*4*+2+/4*-+0*4*01++4*1+-24*+*+.*4*: Occurrence of Leptobryum sp. without moss pillars.: Occurrence of Leptobryum sp. with moss pillars.S. Imura et al.8growing on the ground in Antarctica. There is no apparent terrestrial source popula-tion from which the lake bottom population originated, at least in East Antarctica. Onthe other hand, B. pseudotriquetrum is one of the commonest moss species on the groundin Antarctica. It is thought to be one reason for the di#erence of the distribution ofthese two aquatic mosses in this region.AcknowledgmentsWe wish to express our sincere gratitude to the members of the -0th to .,ndJapanese Antarctic Research Expedition teams for their devoted support to the ﬁeldsurvey in Antarctica. This study was partly supported by a Grant-in Aid for scientiﬁcresearch from the Ministry of Education, Science, Culture, Sports and Technology,Japan (No. +,0.*03+) for S. Imura and H. Kanda and a similar Grant (No. +.-2*,/2)for S. Imura, K. Seto, S. Ohtani, S. Kudoh and H. Kanda.ReferencesArts, T. (,**+): The moss genus Leptobryum and the identity of Pohlia integra. J. Bryol., ,-, -,/--*.Bodin, K. and Nauwerck, A. (+302): Produktionsbiologische Studien u¨ber die Moosvegetation eines KlarenGebirgsees. Schweiz. Z. Hydrol., -*, -+2-/,.Franz, T.C. and Cordone, A.J. (+301): Observations on deepwater plants in Lake Tahoe, California andNevada. Ecology, .2, 1*31+..Ignatov, M.S. and Kurbatova, B. (+33*): A review of deep-water bryophytes with new records from USSR.Hikobia, +*, -3-.*+.Imura, S. and Kanda, H. (+32/): The gemmae of the mosses collected from the Syowa Station area,Antarctica. Mem. Natl Inst. Polar Res., Spec. Issue, .., ,.+,.0.Imura, S., Higuchi, M., Kanda, H. and Iwatsuki, Z. (+33,): Culture of rhizoidal tubers on an aquatic mossin the lakes near the Syowa Station area, Antarctica. Proc. NIPR Symp. Polar Biol., /, +,-+,0.Imura, S., Bando, T., Saito, S., Seto, K. and Kanda, H. (+333): Benthic moss pillars in Antarctic lakes. PolarBiol., ,,, +-1+.*.Kanda, H. and Iwatsuki, Z. (+323): Two aquatic mosses in the lakes near Syowa Station, ContinentalAntarctica. Hikobia, +*, ,3-,31.Kanda, H. and Mochida, Y. (+33,): Aquatic mosses found in lakes of the Skarvsnes region, Syowa Stationarea, Antarctica (Extended Abstract). Proc. NIPR Symp. Polar Biol., /, +11+13.Kanda, H. and Ohtani, S. (+33+): Morphology of the aquatic mosses collected in lake Yukidori, Langhovde,Antarctica. Proc. NIPR Symp. Polar Biol., ., ++.+,,.Kasper, M., Simmons, G.M., Parker, B.C., Seaburg, K.G. and Wharton, R.A. (+32,): Bryum Hedw. collectedfrom lake Vanda, Antarctica. Bryologist, 2/, .,..-*.Light, J. J. and Heywood, R.B. (+31-): Deep-water moss in Antarctic lakes. Nature, ,.,, /-//-0.Light, J. J. and Heywood, R.B. (+31/): Is the vegetation of continental Antarctica predominantly aquatic?Nature, ,/0, +33,**.Light, J. J. and Smith, R.I.L. (+310): Deep-water bryophytes from the highest Scottish lochs. J. Bryol., 3,//0,.Longton, R.E. (+322): The biology of polar bryophytes and lichens. New York, Cambridge University Press,-3+ p.Nakanishi, S. (+311): Ecological studies of the moss and lichen communities in the ice-free areas near SyowaStation, Antarctica. Nankyoku Shiryoˆ (Antarct. Rec.), /3, 0230.Ochi, H. (+313): A taxonomic review of the genus Bryum, Musci in Antarctica. Mem. Natl Inst. Polar Res.,Spec. Issue, ++, 1*2*.Priddle, J. (+313): Morphology and adaptation of aquatic mosses in an Antarctic lake. J. Bryol., +*, /+1/,3.Distribution of aquatic mosses 9Priddle, J. and Dartmall, H.J.G. (+312): The biology of an Antarctic aquatic moss community. FreshwaterBiol., 2, .03.2*.Savich-Lyubitskaya, L.I. and Smirnova, Z.N. (+3/3): New species of Bryum Hedw. From the Bunger Hills.Inf. Byull. Sov. Antarkt. Eksped., 1, -.-3 (in Russian).Savich-Lyubitskaya, L.I. and Smirnova, Z.N. (+30*): New variety of Bryum korotkevicziae Sav. -Lyub. et Z.Smirn. Inf. Byull. Sov. Antarkt. Eksped., +1, ,/,1 (in Russian).Savich-Lyubitskaya, L.I. and Smirnova, Z.N. (+30.): A deep-water member of the genus Plagiothecium Br etSch. in Antarctica. Inf. Byull. Sov. Antarkt. Exped., .3, ---3 (in Russian).Seppelt, R.D. (+32-): The status of the Antarctic moss Bryum korotkevicziae. Lindbergia, 3, ,+,0.Smith, R.I.L. (+32.): Colonization and recovery by cryptogams following recent volcanic activity onDeception Island, South Shetland Islands. Br. Antarct. Surv. Bull., 0,, ,//+.Tewari, S.D. and Pant, G. (+330): Some moss collections from Dakshin Gangotri, Antarctica. Bryol. Times,3+, 1.Wilson, A.T. (+30/): Escape of algae from frozen lakes and ponds. Ecology, .0, -10.(Received July +/, ,**,; Revised manuscript accepted September +2, ,**,)S. Imura et al.10

Distribution of aquatic mosses in the Soya Coast region, East Antarctica

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Distribution of aquatic mosses in the Soya Coast region, East Antarctica

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