The Environment of the Bunger Hills

John A. E. Gibson

Australian Antarctic Division Channel Highway Kingston Tasmania 7050 Australia

September 2000

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Contents

1. Introduction 4 1.1 Geographical setting of the Bunger Hills 4 1.2 Nomenclature 6 1.3 Occupation history 7 2. Weather 13 3. Geology, Geomorphology and Quaternary History 16 4. Lakes 17 4.1 Hydrology 17 4.1.1 Algae Lake drainage system 18 4.1.2 Ice-dammed epiglacial lakes 21 4.1.3 Epishelf lakes 24 4.1.4 Open lakes 25 4.1.5 Closed lakes 27 4.1.6 Ice-based drainage on the Apfel Glacier 28 4.1.7 Water levels and melt intensity 28 4.1.8 Ice cover 30 4.2 Physical limnology 30 4.2.1 White Smoke Lake 31 4.2.2 Lake Pol’anskogo 32 4.2.3 Lake 12 34 4.2.4 Transkriptsii Gulf 35 4.2.5 Lake 14 37 4.2.6 Lake Polest 37 4.2.7 Mixing and history of the epishelf lakes 38 4.3 Chemistry 41 4.4 Biology 45 4.4.1 Non-photosynthetic bacteria 46 4.4.2 Photosynthetic bacteria and algae 46 4.4.3 Fauna 49 4.4.3.1 Rotifers 49 4.4.3.2 Platyhelminthes 50 4.4.3.3 Copepods 50 3

5. Terrestrial Flora 53 5.1 Terrestrial algae 53 5.1.1 Stream algae 53 5.1.2 Epilithic algae 54 5.1.3 Sublithic algae 54 5.1.4 Soil algae 55 5.2 Fungi 55 5.3 Lichens 55 5.4 Mosses 67 5.5 Liverworts 69 5.6 Spatial zonation of flora 69 6 Terrestrial Fauna 70 6.1 Heterotrophic protists 70 6.2 Nematodes 71 6.3 Tardigrades 71 6.4 Mites and fleas 71 6.5 Birds 72 6.5.1 Snow petrel 72 6.5.2 Wilson’s storm petrel 73 6.5.3 South polar skua 73 6.5.4 Adélie penguin 75 6.6 Seals 76 Acknowledgments 78 References 79 Appendices 87 4

1. Introduction

During the 1999/2000 summer a group of four people visited the Bunger Hills (66° 20’S, 100° 50’ E), an area of ice-free rock situated in Wilkes Land, eastern Antarctica, with the aim of gathering scientific and other information to support the development of an Environmental Impact Assessment for the use of sites in the area for landings by large aircraft capable of intercontinental flight. The following report, which describes the natural history of the Bunger Hills, was developed from observations made in the field during this visit supplemented by information published previously.

This report is divided into several sections. The remainder of Section 1 deals with the geographical setting, nomenclature and occupational history of the Bunger Hills; Section 2 covers the weather; Section 3 briefly discusses the geology, geomorphology and Quaternary history; Section 4 covers the hydrology, physics, chemistry and biology of the extensive lake systems of the Hills; Section 5 discusses the terrestrial flora; and Section 6 covers the terrestrial fauna.

1.1 Geographical setting

The Bunger Hills is a rocky, ice-free area located in Wilkes Land, eastern Antarctica. It is approximately 1000 km from Davis in the similarly ice-free Vestfold Hills, 450 km from Casey, and 3800 km from Hobart. It is directly south of the island of Sumatra in Indonesia. The nearest permanently inhabited Antarctic station is the Russian base Mirny, located 340 km to the west, though, as discussed below, three summer stations that are occupied sporadically are located within the Bunger Hills.

The Bunger Hills is surrounded on all sides by ice: to the east lies the Remenchus Glacier and the polar ice cap, to the south the Apfel Glacier, to the west the Apfel, the Edisto Ice Tongue and the Scott Glacier, and to the north the Shackleton Ice Shelf (Figure 1). It is thus different to the Vestfold Hills, which has the open ocean as one of its boundaries, but similar to Schirmacher Oasis in Dronning Maud Land (Bormann and Fritzsche, 1995). The total area is about 950 km 2, of which 420 km 2 is exposed rock (Markov et al., 1970). The remainder is a marine ecosystem that is connected to the Southern Ocean under the Shackleton Ice Shelf, which is a permanent ice shelf held in place by Mill and Bowman Islands. The ice on this marine area breaks out in some summers, and the water, though a little fresher at the surface than seawater, is marine in its 5

(a)

Schirmacher Oasis

Weddell Sea

Beaver Lake Ablation Lake Vestfold Hills Davis South Pole Mirny Bunger Hills

Amundsen Casey Sea

Ross Sea

N Shackleton Ice Shelf

66 ° 00’ S

0 km 10 Remenchus Glacier Currituck Peninsula

Thomas Island

cier Gla disto E Cacopon Inlet ue Tong Scott Glacier Continental Ice Sheet

Transkriptsii Gulf Rybij Khvost Gulf Epishelf lakes Land Algae Lake Other lakes Marine

Lake Ice Pol’anskogo White Smoke 66 ° 30’ S Lake Apfel Glacier 100 ° 30’ E 101 ° 00’ E

Figure 1: (a) A map of Antarctica showing the location of the Bunger Hills and other places mentioned in the text; and (b) a map of the greater Bunger Hills area. 6 chemical and biological characteristics (Kulbe, 1997). The exposed land can be broken up into three main units (Markov et al., 1970): (i) a number of small islands and nunataks in the north of the oasis (Highjump Archipelago); (ii) a series of larger islands and peninsulas located in an east- west band (Geographer’s (Currituck) and Thomas Islands, and Charnockite and Geomorfologov Peninsulas) and (iii) a southern land mass, along with Aviatorov Peninsula and Fuller Island. The last unit was the only one investigated during the 1999/2000 ANARE visit, and, in general, the more northerly units are poorly known. The third unit, which is referred to as the southern Bunger Hills throughout the rest of this report, is approximately 30 km long (east-west) by 20 km wide (north-south), with an area of circa 270 km 2.

The topography of the southern Bunger Hills ranges includes areas of rugged landscape in the south and southeast, with maximum relief of about 160 m. The bedrock, which consists predominantly of granulite-facies orthogneiss cut by alkaline mafic dykes (Sheraton et al., 1997), is dissected by many steep valleys often aligned along obvious linear fault systems (the most prominent of which trend 50°-230°, 120°-300°, and 0°-180°). In many places these valleys are filled by lakes, with the morphology of the largest of these, Algae Lake, which is 14.3 km 2 and 143 m deep, clearly defined by the fault system. To the north and west, the topography flattens out, with isolated hills up to 120 m high separated by areas of till-covered lowlands. Lakes still occur, but are generally smaller and rounder in outline.

1.2 Nomenclature

Various features in the Bunger Hills have been given names by US, USSR, Polish and Australian expeditions. In some cases more than one name exists for the same feature, e.g., Algae Lake (US, and accepted by Australia) compared to Lake Figurno(j)(y)e (Russian). Where relevant, the names and spellings currently accepted by the Australian Antarctic Names and Medals Committee have been used in this report. However, the reader should be aware that, when consulting earlier and/or foreign literature, alternative names and spellings (especially for those transliterated from Russian) might be encountered. All names accepted by the Antarctic Names Committee of Australia in the southern Bunger Hills are shown in Figure 2, along with a number of other unofficial and/or foreign names used in this report.

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Cacopon Inlet

Southern Bunger Hills Gl’atsiologov I. Geologov I. Leon- C. Henderson Fuller I. Edisto Glacier ova Vertoletnyj B. 7 Pen. Magnit I. Magnitologov Str. Chas- Kalach I. ovoj I. C. Aviatorov P. Krylatyj Sur- Pen. Rybij ovyj Krajnij Pen. Korsor I. Shchel’ L. Khvost 3 1 Gulf L. Polest Kinz C. Eolovyj hal B Transkriptsii Gulf Edgeworth Zakrytaja B. David C. T’ulenij C. Hordern Station 9 10 6 8 Soglasija L. Zerkalnoje L. 5 Rogataja Hill Izvilistaja Bay Oasis-2 Dolinnoje L. Station Dobrowolski Station Izumrudn -oye L. L. Dolgoe 4 N Glavnyj I. Dlinnyj Pen. Algae Lake 2 C. Ostryj Vstrech I. L. Burevestnik Tikhoje L. White Smoke L. L. Pt’ich’je L. Pol’anskogo L. Dalekoje

0 5

Apfel Glacier km

Figure 2: A map of the southern portion of the Bunger Hills, showing names accepted by the Antarctic Names Committee of Australia, as well as some unofficial and/or foreign names used in this report (in italics). The map is derived from a Russian map published in the Atlas Antarktiki (1966). Numbers indicate the localities of (1) Granatovaya Sopka Island; (2) Neozhidanniyj Peninsula; (3) Ostrovnaja Bay; (4) Piramidal’naya Hill; (5) Sfinks Hill; (6) Sredn’aja Hill; (7) Storozhevoj Peninsula; (8) Cape Tektonicheskij; (9) Zapadnoye Lake; and (10) Vostochnoye Lake.

1.3 Occupation history

The Bunger Hills were first sighted by a party led by Frank Wild of the Australasian Antarctic Expedition’s Western Party in November 1912. They saw and named Horden Island, now known as Cape Horden, from a distance, but were unable to reach the rocky outcrop due to severe crevassing on the Denman and Scott Glaciers. The full extent of the ice-free area was first recognised in January 1947 by members of Operation Highjump, which had the aim of photographing the Antarctic coastline. A seaplane landing was made in February 1947, and the area was named the Bunger Oasis (later the Bunger Hills) after the pilot of the plane, Lieutenant Commander David Bunger USN. Byrd (1947) described the area as ‘…one of the most remarkable regions on earth. An island suitable for life had been found in a universe of death.’ A further visit took place in January 1948 (Operation Windmill), which provided ground control for the aerial photography taken the previous year.

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The next visit to the area was made by the 1 st Soviet Antarctic Expedition (SAE) from 22- 30 January 1956 (Avysuk et al., 1956; Nudel’man, 1966). Eight scientists were flown to a camp on the ice to the south of the Hills, from where visits were made to ice-free areas in both the Bunger Hills and other nearby outcrops. These initial observations convinced the Soviet authorities to develop a scientific base in the region, even though there were no requirements for it under the plans for the International Geophysical Year (Nudel’man, 1966). The first loads of cargo for the base were flown from Mirny between the 5-13 April. Further equipment was shipped to the Bunger Hills in August, and an unsuccessful search made for a suitable location for a station. Further reconnaissance flights in early September 1956 resulted in the identification of a site next to Algae Lake, with the airstrip situated on the sea ice 3.5 km to the northeast. Workers arrived on 19 September, and construction of the station began. Construction was completed by the middle of October.

The area was also visited in January 1956 by ANARE, though no landings were made. Dr Phillip Law and pilot Doug Leckie visited the Bunger Hills by Beaver aircraft after taking off from sea ice adjacent to the M.V. Kista Dan. The weather closed in, and they returned to the ship after taking a few photographs.

Figure 3: A.B. Dobrowolski Station (originally Oasis Station), January 2000. The partially frozen waters of Algae Lake are visible in the background 9

Oasis (or Oazis) Station was officially inaugurated on 15 October 1956 by the head of the 1st SAE, M.M. Somov (for a photo of the station, see Figure 3). The name of the station recalled the description of Byrd (1947), and highlighted the aims of the SAE in setting it up: to determine why such a large, ice-free area of rock (or ‘oasis’) was present on a continent dominated by ice. Groups of scientists were flown to the Bunger Hills from Mirny for studies of the biology, geology, glaciology and geophysics of the region, and a small group remained over winter to take meteorological and astronomical observations, and to maintain the buildings and equipment. The station was continually occupied by this and subsequent wintering and summering groups until 17 November 1958, when it was temporarily closed down.

At the end of 1958, following a decision of the Soviet government, Oasis Station was transferred to the Polish Academy of Sciences for the continuation of scientific investigations (Nudel’man, 1966). The first group of Polish expeditioners reached Oasis (via Mirny) in mid- January 1959, and the official transfer of the station occurred on 23 January. The station was renamed Antoni Bøleslaw Dobrowolski in honour of a Polish scientist who had wintered with Gerlache’s Belgian expedition of 1897-1899 (Machowski, 1998). The Polish expeditioners left the Bunger Hills after temporarily closing the base on 30 January 1959, planning to return the next summer.

Two Antarctic Historic Sites and Monuments are associated with this early period of inhabitation: Number 10, which is a magnetic observatory with plaque commemorating the opening of Oasis Station; and Number 49, which is a gravity pillar erected by Polish scientists in 1959 (Anonymous, 1997). Both these sites are located within the boundaries of the station.

The next official Polish visit did not occur for another 20 years. In the interim, occasional visits were undoubtedly made by Russian scientists from Mirny, who on at least one, and possibly two, occasions were accompanied by Polish scientists (25 June 1959, though there is some confusion about this date and it is more likely to have been on 25 January 1959, i.e. during the short Polish visit to take over Oasis (Kuc, 1969); 21-22 January, 1966 (Dubrovin and Zalewski, 1969)). The latter is at present the only confirmed visit to the Bunger Hills between 1959 and 1979. Interest in Polish polar research was rekindled in the mid-1970s, which led to a group of scientists visiting from 18 January to the 21 February 1979. Dobrowolski Station was reported to be ‘in good condition’, even though it ‘had not been used for twenty years’ (Krzeminski and Wisniewski, 1985). The work done included geomorphological and geodetic studies, and detailed 10 topographic maps were developed for the immediate area of the station (included as separate sheets in Polish Polar Research, Volume 6: Battke, 1985).

In the meantime, ANARE had taken advantage of an opportunity to visit the Bunger Hills. On 2 March 1977, a party consisting of a biologist, two geologists and a geophysicist were flown into the Bunger Hills by helicopter from the MV Nella Dan, which had reached the edge of the Shackleton Ice Shelf (Barker, 1977). During the short visit to the Bunger Hills, samples of lichens, moss, algae, water and rocks were collected from 14 sites in order to make comparisons with the biology and geology of the Vestfold Hills (see Figure 4 for the location of these sites). An inspection was also made of Dobrowolski Station, which was reported to be in excellent condition considering its age.

This initial Australian visit engendered a deal of scientific interest, and after a few years’ delay a large research program was planned for the 1985/1986 summer. To aid in the planning for this program a second visit by ANARE took place from 6-9 March 1985 (Ledingham, 1985). Two helicopters flew to Dobrowolski from the MV Nella Dan for a short visit. On restart, one of the machines had problems, resulting in spares having to be flown from the ship, and a strip down

Figure 4: Location of sampling sites during the 1977 ANARE visit (Barker, 1977). The numbering system is used throughout this report when referring to samples collected during this visit.

11 of the engine. Thus the trip was extended for longer than anticipated. Advantage was taken of the delay to undertake a small mapping and scientific program. The 1985/1986 summer saw a group of 22 Australian expeditioners in the Bunger Hills (Ledingham, 1986a, b). A new base – Edgeworth David – was established in late January 1986, and officially opened on 8 February (Figure 5). A wide range of scientific studies were undertaken, concentrating especially on geomorphology and geology, with some biology and glaciology. It was anticipated that this season would be the first of three year program, but ice conditions during the subsequent summer precluded further visits, and the program was abandoned. The 1986 visit to the Bunger Hills ended on 4 March.

At the same time, Russian interest in the Bunger Hills was reawakening. Plans were made in 1985 for the construction of a new station in the Bunger Hills (Shabad, 1985), which was built during the 1986/1987 summer, and opened in early 1987 (Filcek and Zielinski, 1990). The station, named Oasis-2, is situated approximately 200 m to the west of Dobrowolski Station (Figure 6). Since the opening of the station, visits were made by the Russians in most summers up until the mid-1990s. A Polish contingent visited in early 1989 (4 February – 10 April), marking the most recent (and longest) visit from that country (Filcek and Zielinski, 1990). A listing of all known visits by Russia and other nations post-1985 is included in Appendix 1, along with

Figure 5: View of Edgeworth David Station, January 2000.

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Figure 6: Oasis-2 Station, located a approximately 200 m to the west of Dobrowolski Station (January 2000). A further building is located close to the edge of Algae Lake. references to the source of information. A number of the Russian visits have involved visiting scientists from American and German institutions.

ANARE visited the Bunger Hills again during the 1995/1996 summer (6 December – 23 February) (Hudspeth, 1996). The party of five people was based at Edgeworth David, but also used field camps throughout the Hills. The majority of the work undertaken was geomorphological in nature, but surveys of flora, fauna and human impacts were also carried out (Hudspeth, 1996; Augustinus et al., 1997; Gore et al., 1999). Two further, short, ANARE visits were made during the 1994/1995 summer to refuel helicopters used in flights between Davis and Casey. The flights refueled at Edgeworth David, and were on the ground for less than 1 hour. A team of five people from the private company Polar Logistics visited the Bunger Hills for ten days in December 1998 while looking for possible blue ice runway sites in eastern Antarctica (M. Sharp, personal communication). This group based themselves at Edgeworth David, but also visited Dobrowolski and Oasis-2 Stations. The 1999/2000 ANARE group of four reached the Bunger Hills by Twin Otter on 26 December 1999, and left on 20 January 2000.

Even though the Bunger Hills has seen sporadic occupation since the late 1950s, there has been a significant human impact on the area. The construction and habitation of the three bases 13 has resulted in considerable amounts of rubbish distributed around the stations themselves, and in plumes to the west blown by the prevailing winds.

2. Weather

One of the major reasons for setting up the original Oasis Station in the Bunger Hills was to determine the climatic conditions which allowed an area of rock to survive in a sea of ice. Thus, the weather records from the early Soviet occupation of the area are remarkably good. These are summarised and interpreted in the Russian literature (e.g. Rusin, 1961), and, fortunately, the data are available in English (Nudel’man, 1966; Gregorczuk, 1980). Table 1 gives a summary of weather data recorded between November 1956 and October 1958. Considerably more data, especially regarding detailed radiation balances, are available in the original publications.

The climate of the Bunger Oasis is similar to that of other rocky coastal areas of the East Antarctic coastline. In most aspects, the climate is close to that of the Vestfold Hills, including

Figure 7: Monthly mean, maximum and minimum temperatures recorded at Oasis station, 1956-8. Also shown (dotted lines) are the equivalent data for Davis station (1957-2000). Table 1: Summary of weather observations for Oasis Station, November 1956-October 1958 (Nudel’man, 1966; Gregorczuk, 1980).

Climate Component Units January February March April May June July August September October November December Year Mean Air Temp. °C 1.8 -2.2 -6 -7.6 -11.3 -20.6 -16.8 -16.5 -16 -11.2 -3.5 1.7 -9.0 Max. Air Temp. °C 10 6 3 1 5 1 -2 -3 -3 0 4 12 12 Min. Air Temp. °C -8 -12 -18 -22 -31 -35 -43 -39 -37 -31 -12 -5 -43 Mean Air Pressure hPa 989.9 988.4 985.4 984 992 997.4 989.2 983.6 980.8 982.6 986 993.6 987.8 Max. Air Pressure hPa 1002.8 1002.8 1004.1 1003.8 1021.1 1027.6 1014.2 1006.7 1001.7 1002.6 997.3 1008.7 1027.6 Min. Air Pressure hPa 975.1 977.2 962.4 937.5 957.4 974.8 952.6 953.7 956.9 968 972.2 978.2 937.5 Mean Wind Velocity m s -1 6.4 5.2 5.1 10 7.6 4.5 7.4 8.6 7.7 5.1 7.7 5.8 6.8 Max. Wind Velocity m s -1 34 34 40 56 40 50 48 49 40 40 45 34 56 No. Days with wind 9 6 9 17 13 7 12 15 13 7 8 6 122 >15 m s -1 Percentage calm % 10.5 13.8 18.2 15.8 25.9 45.2 34 34.4 31.2 26.9 5.0 6.0 22.2 Relative humidity % 45 48 54 53 60 62 70 65 63 56 4 6 48 56 Total Cloudiness % 70 71 69 82 78 60 72 72 71 73 61 62 70 Heavy Overcast % 35 54 42 52 45 22 34 32 44 34 32 3 9 39 Freq. of Cloud % 22 23 26 16 16 34 22 24 24 23 28 30 24 0-20% (total) Freq. of Cloud % 60 40 52 40 45 76 61 64 43 60 62 56 55 0-20% (heavy) Freq. of Cloud % 14 10 9 7 11 14 9 8 8 8 15 15 11 30-70% (total) Freq. of Cloud % 9 15 9 12 16 6 8 6 17 8 9 8 10 30-70% (heavy) Freq. of Cloud % 64 67 65 77 73 52 69 68 68 69 57 55 65 80-100% (total) Freq. of Cloud % 31 46 39 48 40 18 30 30 40 32 28 35 35 80-100% (heavy) No. of fine days (0-20% 4 2 4 2 2 3 2 4 2 4 5 5 39 total cloud) No. of fine days (0-20% 13 5 12 6 8 22 14 14 10 12 11 12 139 heavy cloud) No. of cloudy days (80- 17 14 14 20 18 9 18 17 16 16 12 11 182 100% heavy) No. of cloudy days (80- 4 6 4 6 5 4 4 5 7 4 4 4 57 100% total) Precipitation mm 2.5 3 13.8 4.7 31.3 10.2 17.2 28.8 68.8 13.4 0.5 9.9 204.1 No. days with 6 2 8 6 12 6 10 8 12 9 13 11 93 precipitation >0.1 mm No. days of blizzard 0 2 2 4 6 4 10 8 10 4 2 1 53 temperature regime (slightly cooler at Oasis, especially in winter: Figure 7), pressure, precipitation and cloudiness (Streten, 1986; Streten and Nairn, 1986). It should be noted, however, that the Bunger Hills data were collected over a relatively short period, and longer term averages could smooth out some of the extremes noted. Automatic weather station data collected in 1992-1993 from a site located on an island in White Smoke Lake, close to the southern margin of the Bunger Hills (see Figure 2 for location) (Doran et al., 1996), showed generally similar characteristics to those recorded in the 1950s, suggesting that the data presented here should be accepted as typical for the Bunger Hills. Details of the weather during the 1999/2000 ANARE visit are included in the Field Leader’s report (Kuehn, 2000); in general, the observations from this visit were consistent with previous observations for January.

Plots of wind velocity and direction for Oasis are shown in Figure 8, and indicate that the modal winds are from the east, with a secondary components from the west over the complete year, but with another component from the north, in December. The highest wind velocities were associated with the easterly winds, while the westerly and northerly summer winds were of low velocity. Doran et al. (1996) gave a plot of wind speed versus direction for both winter and summer recorded at White Smoke Lake. Two modal wind directions were again apparent: easterly and westerly. Stronger winds occurred from the east, with speeds up to 18 m s -1 (average 6.2 m s -1), while from the west the average was 2.0 m s -1. Wind speeds from other directions,

(a) (b) 20 20 )

-1 18 18 16 Annual 16 Annual 14 November-February 14 November-February 12 12 December December 10 10 8 8 6 6 4 4 2 2 0 Frequency of Direction (%) Frequency Mean Wind (m Velocity s 0 N NE E SE S SW W NW N N NE E SE S SW W NW N

Direction Direction

Figure 8: (a) Mean wind velocity and (b) frequency of direction recorded at Oasis Station from November 1956 to October 1958 for the entire year, November to February inclusive, and December. Data from Gregorczuk (1980).

16 including the west, rarely exceeded 5 m s -1. These data are consistent with other observations, though the wind speeds seem low (both compared to those given in Gregorczuk (1980) and those observed during the 1999/2000 ANARE visit), perhaps reflecting the relatively protected site of the AWS (especially from winds from the north).

The data set from the 1950s also contains information on cloudiness, which is also summarised in Table 1. This data is a little difficult to interpret, as it is not clear how ‘total cloud’ and ‘heavy cloud’ were defined. However, from the data it is reasonably clear that cloud cover occurs during a significant portion of the year, though a large proportion of this appears to be not included within ‘heavy cloud’. This cloud is likely to be cirrus or similar high cloud.

Observations during the 1999/2000 ANARE visit indicated that there was often a cloud bank directly over the Hills, which presumably formed as a result of warming of moisture-rich surface air by infra-red radiation emitted from the solar-heated dark rock, especially on calm days. This warm air would rise due to its lower density, cool, and eventually become supersaturated with moisture. The sharp margin of this cloud was repeatedly observed to match the edge of the exposed rock, especially along the southern edge of the Bunger Hills. Russian observations suggested that this cloud built up during the morning to a maximum in the middle of the day before dispersing late in the evening as solar heating of the rocks decreased (Gregorczuk, 1980).

3. Geology, Geomorphology and Quaternary History

It is not intended that this report cover in any detail the geology and geomorphology of the Bunger Hills. Articles dealing with these subjects are provided by Adamson and Colhoun (1992), Sheraton et al. (1993, 1995), and Colhoun (1997). Significant work has also been reported on the Quaternary history of the Bunger Hills: Colhoun and Adamson (1989, 1992), Verkulich and Hiller (1994), Melles (1994), Melles et al. (1994, 1997), Kulbe (1997) and Augustinus et al. (1997). Much of this historical information has stemmed from studies of sediment cores collected from lakes or freshwater tidal inlets. A novel approach to the study of the Quaternary history was employed by Verkulich and Hiller (1994), who used radiocarbon dating to measure the age of mumiyo (regurgitated stomach oil of, in this case, snow petrels) in order to determine the age of deglaciation of the Bunger Hills.

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4. Lakes

Lakes and ponds make up an important component of the Bunger Hills landscape and environment. Over two hundred water bodies occur, ranging in size from small, shallow ponds a few tens of metres across that would freeze to the bottom during winter, to Algae Lake, which is one of the largest and deepest freshwater lakes in Antarctica. The salinity of the lakes ranges from ultra-fresh (<20 mg L -1 total dissolved solids (TDS)) for those which receive melt water from the Antarctic plateau, to highly mineralised (>80 g L -1 TDS). In all these aspects, the lakes of the Bunger Hills are similar to those in the Vestfold Hills. A major features that differentiates the lacustrine environments of the southern Bunger Hills, however, is the presence of at least five (and as many as 11 or perhaps more in the greater Bunger Hills area) epishelf lakes. These lakes, which are positioned between land and a floating ice shelf or glacier, typically contain a layer of freshwater overlying water of marine origin, and are tidal. The world inventory of this type of lake, which are thought to be important refugia for the survival of flora and fauna during glacial periods (Bayly and Burton, 1993), consists of only about ten other known examples.

This section discusses various aspects of the limnology of the lakes of the southern Bunger Hills. It includes discussion of previous work (especially on lake chemistry and biology) integrated with observations and results obtained during the 1999/2000 ANARE visit to the area. Four major aspects are considered: hydrology, physics, chemistry and biology.

4.1 Hydrology

Five distinct hydrological regimes were identified in the southern Bunger Hills during the 1999/2000 ANARE visit. Along the southern margin of the Hills, epiglacial lakes (situated between the glacier and rock of the Hills) were identified which drained into Algae Lake, while others either did not have an apparent outlet or were tidal, epishelf lakes. Further epishelf lakes occurred on the western margin of the Hills. Away from the glacial margins, lakes could either be closed (i.e. no outlet) or open, with an outlet either to a closed lake, the ocean, a tidal freshwater epishelf lake, or Algae Lake. A further drainage system occurred on the surface of the Apfel Glacier which resulted in water flowing away from the Bunger Hills.

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Southern Bunger Hills

Sampling site, Transkriptsii Gulf

14

Transkriptsii Gulf

17 13 16 L. Dolgoe N L. Algae 12 15

L. Burevestnik White Smoke L. L. Pt’ich’je L. Pol’anskogo 11 10 9 Lake Dalekoje 8 7 6 1 5 2 4 3 0 5 km

Figure 9: The Algae Lake drainage system. Lakes draining through Algae Lake into Transkriptsii Gulf are shown in dark blue. Also shown are other active or inactive systems noted during the 1999/2000 ANARE visit, and the numbering system used for lakes in this report. It is likely that other lakes close to the eastern ice margin and the northern extensions of Algae Lake also make up part of the drainage system. Also shown is the sampling site in Transkriptsii Gulf.

4.1.1 Algae Lake drainage system

A significant drainage system was identified in the south-eastern section of the study area (Figure 9). The total length of this system, which (in theory) reaches Transkriptsii Gulf at the outflow of Algae Lake, is approximately 25 km long, making it, potentially, one of the longest drainage systems in Antarctica. A similar system that drains a portion of the Sørsdal Glacier occurs in the Vestfold Hills (Bronge, 1996).

The outlet from the westernmost of the epiglacial lakes in this drainage system (Lake 6 in Figure 9) flowed into a small lake (ca. 200 m diameter), which in turn flows into the south western arm of Burevestnik Lake. When visited Burevestnik Lake was not overflowing, but if the water level rose marginally a stream would flow through a snow-filled valley between the lake’s easternmost point and Lake Pt’ich’je. It is likely that this happens in most, but maybe not all, summers. Lake Pt’ich’je, which is at least 22 m deep (though probably much deeper), and had an ice cover 1.5 m thick in January 2000 indicating annual melt out, received water from a number of small lakes abutting the glacier (Lakes 3, 4 and 5) via short streams which run either on the 19 surface of the rock or under snow banks, often via other small lakes. There was also direct input of glacial melt water into Lake Pt’ich’je via a snow bank to the south of its southernmost arm. Lake Pt’ich’je, in turn, flowed from its easternmost point into Lake 2 via a narrow gully. The change in elevation is approximately 5 m. Lake 2 abuts the glacier margin for a considerable distance on its southern side, and thus gets direct melt water input (Figure 10). The outflow of this lake was via a narrow valley to the northeast into Lake 1. The water level drop between these lakes is again small, but a strong stream was flowing when visited on 5 January 2000 (Figure 11), and the lakes should not be considered a single water body as suggested on the Russian map (Atlas Antarktiki, 1966). Lake 1 also abuts the ice margin for a short distance on its southern shore. Its outflow is under a large, permanent snow bank on its eastern shore. The outflow enters a large ice cave, and reappears on the other side of the bank in another ice cave before flowing down a valley to Lake Dalekoje.

Lake Dalekoje also has an ice margin, but in contrast to the other epiglacial lakes, the ice is part of the polar plateau rather than the Apfel Glacier. The lake has markedly different characteristics to the other epiglacial lakes, in that it is highly turbid, resulting from input of glacial flour (very finely ground sediment which remains suspended in the water column) in inflowing water from the base of the ice sheet. In contrast, the lakes described above are

Figure 10: Lake 2, showing the glacial margin forming its southern shore. Looking southeast, 5 January 2000. 20

Figure 11: Outflow from Lake 2 to Lake 1. Looking southwest, 5 January 2000. characterised by extreme clarity of the water. This dichotomy is clearly evident in aerial photographs taken during Operation Highjump in February, 1947 (Byrd, 1947), one of which is reproduced in Figure 12. It thus appears that the input into the drainage system from the Apfel Glacier is largely from surface melt of the glacier rather than basal melting, which possibly reflects the topography of the underlying bedrock (i.e. sloping away from the southern margin of the Bunger Hills). An area of non-turbid water was clearly apparent on 5 January 2000 at the inlet of the stream from Lake 6 into Lake Dalekoje. Lake Dalekoje, which is at least 40 m deep (Klokov et al., 1990), drains via a long, narrow, snow-filled valley that extends from the western extremity of the lake to the narrow southeastern arm of Algae Lake. No convincing evidence of flow between the lakes was observed during the 1999/2000 ANARE visit, though an under ice stream was observed approximately halfway along the valley. Evidence of flow from Lake Dalekoje to Algae Lake stems from Figure 12, which clearly shows a melt stream between the lakes, and, importantly, the presence of glacial flour in the arm of Algae Lake into which the stream flows that could only have come from Lake Dalekoje. The other lakes in the drainage system along the margin of the Apfel Glacier are clear in this photo, indicating the absence of input of glacial flour even during this period of apparently higher melt.

21

Figure 12: Photo reproduced from Byrd (1947) of a portion of the Algae Lake drainage system. Of particular note are clear, ice-free lakes abutting the Apfel Glacier (Lakes 1 and 2, with Lake Pt’ich’je just visible at the right hand margin), and the presence of glacial flour in Lake Dalekoje and the south-eastern arm of Algae Lake.

Algae Lake is a large (14.3 km 2), deep (143 m) lake whose alignment is defined by clearly marked fault lines which traverse the southern Bunger Hills. The lake also abuts the glacial margin for circa 400 m at its eastern extremity, and receives direct and other, indirect, glacial melt water inputs. These sources were not investigated during the 1999/2000 ANARE visit. Water from the drainage system eventually exits Algae Lake via an outlet stream that flows to Transkriptsii Gulf from the lake’s westernmost point. No water flow was observed in this stream during the 1999/2000 ANARE visit, but it flowed the previous summer (M. Sharp, personal communication) and in other years (Barker, 1977; 1986 ANARE aerial photography; Klokov et al., 1990).

4.1.2 Ice-dammed epiglacial lakes

Epiglacial lakes abut both glaciers and rocks for portions of their shoreline. This lake type is widespread in Antarctica, as they occur around the margins of many nunataks and oases. The 22 epiglacial lakes along the southern margin of the Bunger Hills either flow out to the drainage system described in Section 4.1.1, or are ice-dammed. None of the ice-dammed lakes currently have outlets, but if they did, they would flow over ice (except for Lake 8; see below) to topographic lows along the ice-rock margin of the Hills. These lakes are described in this section.

Water was not flowing out of the next major lake to the west of Lake 6 (Lake 8, Figure 9) when visited (7 January 2000), but if the water level were to do so (it would have only required a slight increase in water level for flow to occur when visited) it would clearly flow to a small lake to the northwest. This lake in turn would overflow through a narrow gully into a topographical low at the southern end of a major snow bank. This low was occupied by a small area of flat, clear lake ice (as distinct from the blue, bubble-filled glacier ice that surrounded it) (Lake 9), but the low water/ice level indicated that little if any melt reached this area. No attempt was made to drill a hole through the ice of Lake 9 to determine if unfrozen water was present. The logical outflow from Lake 8 might appear to be through a series of medium sized lakes into Lake Burevestnik, but this route is blocked by a moraine ridge approximately 5 m above lake level. Lake 8 receive melt water input directly from the glacier, as well as possibly from a small lake (Lake 7) also abutting the glacier to the south-east. This lake was completely ice-covered when visited and would overflow into Lake 8 if water level increased. In this region the margin of the glacier was not characterised by cliffs (as it was for most of the lakes to the east), but rather sloped down more gently on ice and snow banks to the lakes.

The next lake to the west of Lakes 8 and 9 was White Smoke Lake. The name of this lake, which was coined during the joint US-Russian visit in 1991/1992, is not recognised by either the Australian or US names committee, but is in common usage in the literature (e.g. Bayly, 1994; Doran et al., 1996, 2000). This large, roughly rectangular lake, circa 2 km long and 500 m across, is bounded on its eastern, northern and western margins by rock or permanent snow banks, and by a steeply but smoothly sloping ice ridge to the south. Over the ridge the ice slopes down to the south to a topographic low to the west (i.e. downstream in a glacial sense) of an isolated nunatak. The ridge is not as marked at its western end, and if the lake were to overflow it would be via this gap to Lake 10 (see below). However, the gap is currently the main route by which melt water reaches the lake from the glacier; on the 13 January 2000, the ice in this gap was very slushy and water was flowing into the lake. Some melt will also come from the land margins. White Smoke Lake is at least 90 m deep (Bayly, 1994), and is of very low salinity (see Section 4.2.1), reflecting the major water input being from glacial melt. A relatively large 23

‘iceberg’ was observed in the lake to the west of a small island in the centre of the lake. Tide gauge data collected in 1991/1992 indicated that White Smoke Lake was tidal (Bayly, 1994; Doran et al., 2000), with a range of circa 30 cm. Tide cracks are clearly evident in photographs taken at this time, and water was recorded on the surface of the ice (D. Andersen, personal communication). No evidence of tidal action was observed during the 1999/2000 ANARE visit suggesting that glacier movement between 1992 and 2000 had severed the hydraulic connection with marine water.

To the west of White Smoke Lake was another topographic low (Lake 10); if White Smoke Lake were to overflow, water would flow over the ice surface and reach this basin. This area was covered by snow when visited, but an excavation at its lowest point revealed clear lake, rather than bubbly glacier, ice (Figure 13). Water would also reach this basin from Lake 11 to the west. This lake, which was considerably smaller than White Smoke Lake, was again bounded by glacial ice on its southern margin, and by land on its other sides. The lake ice was close to the level required for outflow, which would be to the east through a gap in the rocks to Lake 10. No evidence of flow was observed, however. The surface level of the ice was estimated to be 20 m above sea level. The ice on this lake was at least 4 m thick (the maximum length of the ice augers

Figure 13: Lake ice under snow in the depression to the west of White Smoke Lake (Lake 10). 24 available), and, in contrast to the hole in White Smoke Lake, no water accumulation in the hole in the ice from subsurface melt. It is possible that Lake 11 was frozen to its bottom.

4.1.3 Epishelf lakes

Epishelf lakes are similar to epiglacial lakes in that they abut both ice and land, but in this case the ice is a floating ice shelf or glacier. There is thus the possibility of hydraulic connection with marine water under the ice, and such lakes are tidal. As discussed above, there are only very few epishelf lakes known, and a high percentage of these occur in the Bunger Hills.

Lake Pol’anskogo, the next lake to the west of Lake 11, has been reported to be tidal by many workers (e.g., Gal’chenko, 1994; Melles, 1994). When visited on 13 January 2000, the presence of large tide cracks confirmed this lake’s inclusion in the inventory of epishelf lakes. Like White Smoke Lake, Lake Pol’anskogo is bounded to the south by ice, but in this case heavily eroded and crevassed ice cliffs are present. Rock and permanent snow banks surround the lake on its other sides, and a number of small islands is present. The tidal range is probably similar to that observed in Transkriptsii Gulf further to the north. The lake therefore has an hydraulic connection under the ice of the Apfel Glacier. It has two major basins, one of which contains cold freshwater and the other relatively warm saline water (see Section 4.2.2). Water input into this lake will also occur from a limited land-based drainage area, including a small lake on the rock to the east.

From Lake Pol’anskogo the margin of the southern Bunger Hills is generally aligned north-south, then northeast-southwest. All the lakes along this margin are tidal, and their physical characteristics are discussed in more detail below (Section 4.2.3-5). The first of these (Lake 12) has depth of at least 65 m, and contains brackish water overlying more saline water. A very small tidal lake (Lake 13), just visible on the ANARE satellite image map but clearly evident in the 1986 ANARE aerial photography, occurs between Lake 12 and Transkriptsii Gulf on the landward side of the moraine line. It is possible that this moraine is ice-cored, and the tidal connection occurs through cracks in the ice. Transkriptsii Gulf (Figure 14) is the largest of the tidal lakes, being some 8 km long and 1-2 km wide, and bounded by land to the east and, except 25

Figure 14: View across Transkriptsii Gulf from Edgeworth David Station in the late evening. January 2000. The land to the right is Krajnij Peninsula. for two ‘peninsulas’, ice to the west. The maximum depth of the Gulf is over 120 m (Klokov et al., 1990). Transkriptsii Gulf has inputs of freshwater from Algae Lake, as well as short outflow streams from other lakes. An extensive drainage system also flows into the lake from the Scott and Apfel Glaciers; this drainage is mapped on the Russian maps, and is clearly visible on the ANARE satellite image map.

The northernmost tidal lake (Lake 14: Figure 15) is much smaller than Transkriptsii Gulf, and is at least 22 m deep. Some inflow into the lake would occur from local lakes as well as melt from the surface of the ice sheet. This lake is north of the Apfel-Edisto medial moraine, and therefore is bounded to the west by the Edisto Ice Tongue rather than the Apfel Glacier. The lake has contact with the ice to the west through a break in the Younger Edisto Moraine.

4.1.4 Open lakes

Open lakes are those which have an outflow. The presence or absence of an outlet is not absolute; water levels can fall, resulting in loss of outflow, or rise, causing a previously closed

26

Figure 15. Aerial view of Lake 14 (taken during the 1985/1986 ANARE visit), showing the proposed boundary between the basins resulting from the Edisto Moraine continuing under the surface of the lake, and the approximate position of the sampling site. lakes to overflow. The former process was clearly at play in the Bunger Hills during the 1999/2000 ANARE visit.

Open lake systems generally occurred in the southern portion of the southern Bunger Hills to the south of Algae Lake (Figure 9). There appeared to be greater accumulation of snow melt into the drainage basins of these lakes, and therefore water input into the lakes. Strong drainage systems (in addition to those discussed above) were observed, for example, between Lakes 15-17, eventually flowing into Algae Lake. Other systems were clearly evident either from the aerial or satellite photos, but were largely inactive during the 1999/2000 ANARE visit. For example, Lake 15 overflows via Lake 16 to Lake Dolgoe, with water flowing from this lake near its eastern extremity into Algae Lake. However, there was no flow into Lake 16, or out of this lake to Lake Dolgoe on 7 January 2000. A few days later the outflow from Lake Dolgoe was visited, and no flow was occurring. In contrast, aerial photography taken in 1986 showed these streams flowing strongly, indicating that the 1999/2000 summer was one of low melt. Other evidence supporting this conclusion included the presence of algal stains well above the current level of Lake 16 (Figure 16), indicating low water level.

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Figure 16: Algal stains on rocks at the edge of Lake 16, indicating low water level. The stains extend for approximately 30 cm above current water level.

In general, most of the larger lakes to the south of Algae Lake and Transkriptsii Gulf appeared to be part of drainage systems that may have been inactive during the 1999/2000 summer, but which would have been active during summers of greater melt. The streams in these drainage systems were particularly important in the botany of the regions, often having rich moss and lichen communities. Very few saline lakes were noted in this region, and these were generally small ponds in fully enclosed basins.

4.1.5 Closed Lakes

Closed lakes dominated the landscape to the north of Algae Lake. These lakes were generally in bowl-shaped depressions covered by a thick layer of glacial till (Figure 17). This was in contrast to the lakes to the south of Algae Lake, which often occurred at the base of steep sided, well- defined valleys. The closed lakes were typically quite saline, with the maximum salinity recorded being over 80 g L -1 (see Section 4.3 below). The water levels in most of the closed lakes was low, as indicated by extensive areas of dried microbial mat around their margins. The margins of the lakes also often had extensive surficial salt deposits. These observations did not preclude the occurrence of open, relatively fresh lakes in the northern part of the study region. For example, drainage systems were noted in a rugged area of Vertoletnyj Peninsula (Figure 9).

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Figure 17: Lake Polest, a closed lake to the north of Edgeworth David Station. Note the presence of thick and extensive deposits of glacial till in the valleys surrounding the lake. View is to the east.

4.1.6 Ice-based drainage on the Apfel Glacier

Most of the surface of the Apfel Glacier did not drain directly into the southern Bunger Hills, but rather surface water flowed in supraglacial streams to the south and the west. These flows eventually reached lakes on the southern margin of the Apfel Glacier near the medial moraine line with the Scott/Edisto Glaciers (Figure 18). Some water also reached these lakes in streams crossing the moraine line from the Scott Glacier. Water in these supraglacial lakes could eventually reach Transkriptsii Gulf. A well-developed stream line is evident both on Russian maps (Atlas Antarktiki, 1966) and the ANARE satellite images of the area flowing on the surface of the Apfel Glacier from the south west into Transkriptsii Gulf. The lie of the land suggests that the underlying bedrock slopes away from the southern margin of the Bunger Hills, and it is probable drainage on the surface and at the base of the glacier is controlled by the underlying topography.

4.1.7 Water levels and melt intensity

A consistent observation during the 1999/2000 ANARE visit was that water levels in most, if not all, of the closed lakes (i.e. without stream input and output) was very low. This was indicated by the large rings of dried microbial mat material around the margins, increased salinity 29

Figure 18: Drainage channels and supraglacial lakes on the surface of the Apfel Glacier to the south of the Bunger Hills. The medial moraine between the Scott and the Apfel Glaciers is visible in the centre of the image. The view is looking to the east southeast .

in Lake Polest (see below), and geomorphological evidence such as inactive erosional gullies and streams. It is not certain whether this was a long term trend (i.e. these gullies have been inactive for many years), or whether this reflected conditions during this summer only. The 1999/2000 summer was one of the coldest on record at Davis and Casey (J. Jacka, personal communication), suggesting that the intensity of the melt during the visit to the Bunger Hills could have been lower than in most years. The low lake levels reflect low input of water from snow banks and other sources in the drainage basins. This in turn may indicate reduced accumulation of precipitation or reduced melt due to cooler conditions. Support of the latter conclusion also came from the relatively low melt water flow from the Apfel Glacier into the drainage system described above. Though there is no evidence of what is ‘normal’, flow out of Lake Pt’ich’je, which should have integrated melt from a large area of the glacial margin, was only of the order of tens of litres per minute. This compares with systems such as Ellis Rapids in the Vestfold Hills, which has many orders of magnitude higher flow in summer (Bronge, 1996). The closed lakes of the Vestfold Hills are currently going through a period of decreased water level (Gibson, 1999), and it may be that this trend is spread at least along this section of the coast of East Antarctica. 30

4.1.8 Ice cover

All of the lakes of the Bunger Hills were ice covered in late December at the start of the 1999/2000 ANARE visit, though many had a small, open area (in most cases less than 1% of the total area of the lake). The only lakes to melt out completely during the visit (i.e. up to late January) were small saline lakes to the east and north of Edgeworth David Station, with larger saline (e.g., Lake Polest) and freshwater lakes retaining most of their ice cover until at least the end of January. The only other lakes to show significant ice melt were those in the Algae Lake drainage system through which melt water passed. This level of ice melt appeared to be much lower than typical for this time of year. The Operation Highjump and ANARE aerial photos taken in 1947 and 1986 respectively, as well as the ANARE satellite image map, show most of the lakes except for White Smoke Lake and the other ice-dammed epiglacial lakes, and the epishelf lakes ice-free. The cold summer had presumably resulted in delayed melting of the ice.

Ice thickness on the permanently ice covered epishelf lakes appeared to have increased significantly between 1991/1992 (Gal’chenko, 1994; Doran et al., 2000) and 1999/2000. Table 2 gives a comparison of recorded thicknesses; these data suggest a cooling of the climate of the area over this period. Further monitoring is required to determine which climatic factors are responsible for controlling ice thickness.

Table 2. Ice thickness (m) on selected lakes measured during the summer of 1991/1992 (D. Andersen, personal communication) and 1999/2000.

Lake Ice Thickness 1991/1992 1999/2000 Transkriptsii Gulf 2.6 m 3.9 m Pol’anskogo Lake 2.2 m 3.5 m White Smoke Lake 2.3 m 3.75 m

4.2 Physical limnology

Little information has been published regarding the physical structures of the lakes of the Bunger Hills. Algae Lake is known to be over 140 m deep, and very limited data indicate that its 31 temperature remains in the range 0-5 °C for the entire year (Korotkevich, 1964c; Grigor’ev, 1964; Gal’chenko et al., 1995). This thermal regime is typical for a deep fresh water body in polar regions, and is similar to that of Crooked Lake in the Vestfold Hills, which is comparable in maximum depth and area (Laybourn-Parry et al., 1995). Limited salinity data has been reported for Transkriptsii Gulf (Klokov et al., 1990; Gal’chenko et al., 1995), which indicate the presence of a layer of water of approximately seawater salinity between a depth of 88 m and the bottom of the lake at circa 120 m. Data from Lake Pol’anskogo (Gal’chenko et al., 1995) indicate a novel structure, in which one basin is warm and brackish, the other cold and fresh. Finally, data from a few depths have been published for a series of smaller lakes, including Lakes Dolgoe, Dolinnoje, Dalekoje, Polest, Zapadnoye and Vostochnoye (Kaup et al., 1993), an unnamed lake near Oasis Station (Markov et al, 1970), and various lakes between Edgeworth David, Oasis-2 and White Smoke Lake (Gal’chenko et al., 1995). Little can be deduced from these data regarding the structures of these lakes except for the possibility of meromixis (permanent stratification) in Lake Polest, and that the others appeared to be mixed.

Detailed salinity-temperature-depth profiles of six lakes were recorded using a Falmouth Instruments conductivity-temperature-depth (CTD) unit during the 1999/2000 ANARE visit. These profiles, along with other aspects of the physical limnology of the lakes, area discussed in detail in the following sections.

4.2.1 White Smoke Lake

Temperature and salinity profiles of White Smoke Lake were recorded on 13 January 2000. The ice thickness at the profiling site was 3.75 m, and the ice cover is permanent (i.e there is no evidence of it melting since the 1950s). The maximum depth of the lake is at least 90 m (Doran et al., 2000), but at the profiling site was only 52 m. The profiles (Figure 19) indicated that the lake was nearly isothermal (at circa 0.33-0.34 °C) and isohaline (at circa 0.035 g L -1) over most of the profile. Cooler water was present close to the surface where it was in contact with the ice and therefore must be at its freezing point, and slightly warmer water was present close to the sediment. A slight maximum in salinity was apparent at about 10 m. The temperature and salinity profiles were similar to those recorded in 1991/19922 (Doran et al., 2000). The earlier profiles, however, extended deeper into the water column, and indicated the presence of slightly more saline and cooler water from 58 m to the sediment. These observations are indications of the tidal nature of the lake at that time. 32

Figure 19: Temperature and salinity profiles of White Smoke Lake, 13 January 2000.

The water salinity as calculated by the CTD was very low, and the variations observed may have been to some extent the result of problems with the measurements of such low salinities by a unit that incorporated software and hardware largely designed for use in more saline water. However, the salinities were similar to those recorded in a previous study for other lakes in the Bunger Hills which receive melt water from Antarctic plateau (Klokov et al., 1990; Doran et al., 2000). This low salinity suggests little if any mixing with marine water; Lake Pol’anskogo, which should receive most of its water from a similar source to White Smoke Lake, but which definitely has contact with marine water, was approximately ten times more saline.

The cold temperature of the lake, only marginally above freezing, is not surprising given the permanent ice cover and the ice margin to the lake. Any heat input into the lake by solar radiation or other sources will result in either the ice cover or the ice margin melting. The temperature of any stream flow from the glacier is likely to be very close to its freezing point, so minimal heat will be imported into the lake by this route. Little is known about horizontal mixing in a water body such as White Smoke Lake, but it is likely that there is a temperature gradient between the (colder) ice margin and the (warmer) rock margin, which will engender mixing in the lake. Without further profiling it is difficult to estimate the intensity of this gradient.

4.2.2 Lake Pol’anskogo

Previous studies in Lake Pol’anskogo indicate that the lake is tidal, and that it is divided into two basins, one of which contains freshwater and the other brackish (Gal’chenko et al., 1995). This 33 structure is possibly unique. The thickness of the ice when sampled on the 13 January 2000 was 3.5 m (c.f. 2.2 m; Gal’chenko et al., 1995), and the water depth 68 m (c.f. maximum recorded by Gal’chenko et al. (1995): 69 m). The salinity and temperature profiles (Figure 20) indicated that the limnology of the freshwater basin was similar to White Smoke Lake (at least to the depth of the profiles) in that the temperature was close to freezing throughout the water column, and that there was little structure except for cooler water immediately under the ice and a slight temperature maximum at about 5 m. This region of higher temperature would be expected to lead to slight instability in the water column, as at these temperatures, warmer water is denser. This apparent instability might have been compensated for by slightly lower salinities, but the change in salinity required was probably beneath the accuracy of CTD unit.

Figure 20: Temperature and salinity profiles of Lake Pol’anskogo, 13 January 2000.

The salinity of the lake was for most of the profile approximately ten times that of White Smoke Lake, though with a fresher layer near the surface. The salinity was also considerably above that expected for a lake that received most of its water input from glacier ice, though the fresher layer at the surface may indicate such an input. As Lake Pol’anskogo is tidal, there must be some hydraulic connection to the ocean. The processes which lead to the contrasting basins of the lake are discussed further below.

The salinity and temperature profiles of the saline basin recorded by Gal’chenko (1995) were very different. The profiles were very similar to that of the freshwater basin to a depth of 28 m, beneath which both temperature and salinity increased to maxima of 5.7 °C and 5.5 g L -1.

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4.2.3 Lake 12

Temperature and salinity profiles of Lake 12, which was also clearly tidal, were recorded on 8 January 2000 (Figure 21). The profiles were more complex than those recorded for White Smoke and Pol’anskogo Lakes, with significantly more saline water present at the bottom of the profile. It is probable that deeper sites with higher salinity occurred somewhere within the lake (c.f. Transkriptsii Gulf); no attempt was made to find the deepest point of the lake due to the presence of a 3.8 m thick ice cover.

Figure 21: Temperature and salinity profiles of Lake 12, 8 January 2000. The dotted line in the temperature plot is the calculated freezing point of the water, and the two numbers on the salinity plot are the salinities in the near isohaline intervals.

The salinity of the water just beneath the ice was 0.59 g L -1, which was slightly more saline than the water in Lake Pol’anskogo, and at about 25 m increased to 1.09 g L -1. This salinity was very similar to that recorded at intermediate depths in Transkriptsii Gulf and Lake 14, and has implications for the origin and mixing of these epishelf lakes (discussed further below). The salinity increased again from a depth of 52 m to the bottom of the profile. The maximum salinity, about 4.5 g L -1, is slightly more than 10 % of seawater salinity. The salinity curve showed a few very small jumps, but the increase was generally smooth. It is not determined if this more saline water was anoxic.

The temperature profile indicated that some near surface heating had occurred (to a maximum of 0.57 °C) at a depth of 15 m. It is possible that this maximum reflected warming at some stage when the ice was thinner and more solar radiation could reach the water column, as no surface warming was evident at the top of the profile. Another source of this heat could have been 35 in freshwater entering the lake from the land. Beneath the temperature maximum, the water cooled, and for most of the section of intermediate salinity the water was only slightly warmer that its freezing point. As salinity at the base of the profile increased, the temperature fell, and approached very closely the calculated freezing temperature. This temperature profile, with most of the water close to its freezing point, is very unusual. Most Antarctic stratified lakes contain warmer water beneath a cool surface layer (Gibson, 1999). It appears in Lake 14 that heat has been removed from the bottom layers until the point has been reached that the water is at its freezing point, and any further heat loss to a heat sink (e.g. seawater at its freezing point (-1.87 °C) or ice) will be accompanied by ice formation.

4.2.4 Transkriptsii Gulf

Transkriptsii Gulf is the largest of the epishelf lakes in the southern Bunger Hills, measuring approximately 8 by 2 km. A long, sinuous inlet (Izvilistaja Bay) is located on the eastern side of the Gulf, and it is at the end of this inlet that the major terrestrial freshwater input occurs (Figure 9). This bay is in places quite shallow, with rock bars exposed at low tides. Break out of the ice occurs in this region, but the remainder of Transkriptsii Gulf was covered by a layer of ice that was 3.9 m thick at three different sites when drilled during the 1999/2000 ANARE visit. The maximum reported depth of Transkriptsii Gulf is 122 m (Melles, 1994), with the deepest point located in the northern section off Krajnij Peninsula. The ice margin of the Gulf appears to be relatively consistent in its position (by comparison to Russian maps derived from aerial photography from the 1950s and 1960, and the ANARE satellite image map (image recorded in 1990), and it is probable that extensive freshwater reaches the Gulf from the ice, especially along a well-defined drainage channel that enters the Gulf to the north of a small peninsula west of Cape Hordern. The peninsulas mentioned in this paragraph are positioned between the ice of the Gulf and the glacial margin, and possibly should be thought of as islands. Three other true islands are located near the entrance to Izvilistaja Bay.

Temperature and salinity profiles were recorded for Transkriptsii Gulf at a site (66° 14.376’S, 100°, 35.426’ E: see Figure 9) reasonably close to the deepest point. Repeated attempts to record a profile closer to the deepest point failed due to instrumental problems. The profiles, which reached a depth of 98 m, indicated that over 80 m of slightly brackish water overlay more saline water, with the maximum salinity recorded approaching that of seawater (Figure 22). The salinity in the brackish zone was 1.05 g L -1, similar to that in Lakes 12 and 14. The salinity 36

Figure 22: Temperature and salinity profiles of Transkriptsii Gulf, 11 January 2000. The dotted line in the temperature plot is the calculated freezing point of the water, and the number on the right hand plot is the salinity in the isohaline interval. increase beneath 80 m was quite smooth, though some structure was evident in the profile. The profile was consistent with earlier Russian work (Klokov et al., 1990), which reported a midwater salinity of circa 1 g L -1, and anoxic saline water beneath a depth of 88 m. From the profiles given here, this interface appears to occur at the first significant increase in salinity above 1.05 g L -1, which is also consistent with chemical data given by Klokov et al. (1990). The tidal range in Transkriptsii Gulf is approximately 2 m, so the actual depth of this interface relative to the ice surface will depend on the state of the tide. The input of freshwater into the Gulf will also affect the position of this interface – this is discussed further below.

The temperature profile of Transkriptsii Gulf was more complex than for the other epishelf lakes, though it was again characterised by very cold water temperatures only slightly above freezing. Even though none of the water was at its freezing point, at the base of the brackish zone it was only marginally warmer. The cooler water near the surface is underlain by a zone, which extends to nearly 40 m, where temperature is reasonably constant at circa 0.25 °C. Beneath this zone complex thermal behaviour occurs, with small peaks and troughs superimposed on a general decrease in temperature with depth. The saline water at the bottom of the Gulf was associated with a marked, but in absolute terms only slight, increase in temperature.

37

4.2.5 Lake 14

The profiles of Lake 14, recorded on 9 January 2000, exhibited features quite different to the other epishelf lakes (Figure 23). Firstly, the lake (at least at the sampling point) was much shallower (22 m). Considering the less rugged topography of the area, this is not too surprising. Secondly, the lake was far warmer (maximum: 5.2 °C), with the temperature in the water column near freezing only directly under the ice. The temperature increase apparent beneath 8 m did not coincide with an increase in salinity, indicating that there was little vertical mixing in the near isohaline water that extended from 3 m to 12 m. Thirdly, salinity increased at a much shallower depth than in Transkriptsii Gulf or Lake 14. As the profiling point was unlikely to be the deepest point in the lake, it is probable that higher salinities, possibly reaching that of seawater, occurred at depths below 22 m. Attempts to determine if the more saline water was anoxic were inconclusive. Even though high salinity water was present, there was still an interval of water (from just under the ice to about 12 m) that had a very similar salinity (1.03 g L -1) to that of mid- water depths in Transkriptsii Gulf and Lake 14.

Figure 23: Temperature and salinity profiles of Lake 14, 9 January 2000. The number on the right hand plot is the salinity of the water in the isohaline interval.

4.2.6 Lake Polest

Lake Polest, which is perhaps the most studied of the saline lakes of the Bunger Hills, is located (Figure 2) to the north of Edgeworth David Station between Transkriptsii Gulf and Rybij Khvost Gulf. Earlier work (Kaup et al., 1993; E. Kaup, personal communication) suggested that the lake may be meromictic, and therefore similar to many lakes in the Vestfold Hills (Gibson, 1999). To determine if stratification existed, salinity and temperature profiles were recorded on 38

11 January 2000. The lake was found to be close to isothermal and isohaline except for a small region immediately under the ice, which would have resulted from summer melt of the fresher ice cover. The salinity of the water, 81 g L -1, was slightly higher than the maximum recorded in the lake in 1989 (79 g L -1), when some stratification, indicated by lower surface salinities, occurred in the surface water (Kaup et al., 1993). The temperature throughout the water column in 2000, -0.5 °C, was far colder than in 1989, when the temperature in the bottom water reached 17.7 °C.

It would appear that a reduction in water level has lead to an increase in salinity in the lake since 1989, and overturn. Such a reduction in water level is consistent with observations made at other lakes in the area during the 1999/2000 ANARE visit. The temperature decrease is also consistent with whole lake mixing. However, after complete melt out of this ice, a layer of less dense water would cap the lake providing conditions for heating of the subsurface water, and it is possible that water temperature reached levels similar to those recorded in 1989 later in the summer. It is probable that no meromictic lakes occur in the Bunger Hills, with the exception of the four stratified epishelf lakes discussed above.

4.2.7 Mixing and history of the epishelf lakes

The profiles of the epishelf lakes presented above provide insight into their mixing and history that complements information available from sediment cores and other sources. In particular, consideration of the physical structure of the lakes allows a better understanding of the processes, probably related to changes in freshwater input, which led to the invasion of seawater into Transkriptsii Gulf, Lake Pol’anskogo and Lakes 12 and 14 on a number of occasions during the Holocene (Melles, 1994; Kulbe, 1997).

The processes occurring in Transkriptsii Gulf are the key to understanding the effects of changing freshwater input. The position of the saline-fresh interface in this water body will be a function of the freshwater balance at the surface. This balance is the net sum of inputs from the Algae Lake (and other terrestrial drainage systems) and the drainage from the Apfel Glacier, and losses from ablation of surface ice. During periods of negative water balance, the depth of the interface will be closer to the surface due to reduced hydrostatic force of the freshwater layer (Figure 24). In contrast, positive water balance will depress the interface, eventually leading to the point where it occurs somewhere under the floating glacier ice in the connection between the Gulf and open ocean, which appears to be the situation at present. The anoxic conditions beneath 39

Figure 24: (a) Conjectured current situation in Transkriptsii Gulf derived from the salinity and temperature profiles given in Figure 21; (b) Expected changes in Transkriptsii Gulf if input of freshwater from melt and runoff were reduced.

88 m (Klokov et al., 1990) suggest that little mixing is occurring in this water, and that the connection between the Gulf and open ocean occurs at a depth above 88 m.

The location of the connection between the marine water and Transkriptsii Gulf is not clear. The occurrence of two ‘peninsulas’ breaking up the ice margin of the Gulf indicates that it cannot be along the entire western margin. The considerable tidal range in the Gulf, especially taking into account its large area, argues for a connection of large cross section. The temperature minimum near 80 m probably reflects the depth of the connection; this is consistent with the freeboard of ice (5-10 m) on the western margin of the Gulf. The freshwater water that is present under the ice will be cooled both by contact with the ice and cold seawater in the under-ice connection to near freezing. That it is not seawater that is currently moved tidally into and out of the lake is indicated by the anoxia beneath the depth at which the salinity begins to rise (at 88 m in the profile of Klokov et al., 1990): if seawater were to enter the Gulf, it would probably be oxygenated, and would flow into the depression beneath 88 m at the base of the Gulf. At present, on entering the Gulf the cold freshwater from beneath the ice would rise, as it is less dense than the slightly warmer water above it (the temperature of maximum density for water of salinity 1 g L-1 is circa 3.8 °C). The peaks and troughs present in the temperature profile indicate that this mixing is a complex process. The heat in the warmer water at the surface will be due to input of warmer water (up to 5 °C) from the Algae Lake drainage basin, and, to a limited extent, solar radiation.

If freshwater input to Transkriptsii Gulf is reduced, the interface between fresh and saline water, currently somewhere under the ice shelf, would reach the lake. This water would initially 40 flow into the deep basin beneath the connection as mentioned above, but, on continuing negative water balance, begin to fill the lake from the bottom. In the extreme case (i.e. if there were limited freshwater input for an extended period), marine water would eventually fill the basin to the top. There is some evidence that this has happened in the past: Verkulich and Melles (1992) and Melles (1994) retrieved sediment cores from Izvilistaja Bay which included at least three marine horizons, indicating invasion of seawater into Transkriptsii Gulf sometime during the Holocene, and sub-fossil sponges were recovered from a tidally-pushed beach ridge near Cape Hordern during the 1999/2000 ANARE visit. By dating periods of marine influx, estimates of the timing of dry (=cold) periods in Antarctica could be determined. The situation would then be like the ‘marine water’ that is present in the central part of the greater Bunger Hills, which can also be considered an epishelf lake as there is a connection with the open ocean under the Shackleton Ice Shelf. In this case, however, freshwater input has not been sufficient to create a markedly lower salinity layer at the surface, and probably has never done so. This region currently possesses a typical marine fauna, though with the possibility of unique species (Brodsky and Zvereva, 1976; Melles, 1994)

The profiles of the other epishelf lakes also provide significant information about processes occurring in the system. In Lake Pol’anskogo, the near constant salinity and temperature argues for a connection to marine waters well beneath the maximum depth of the profile, that the fresh-saline interface is well away from the lake, or that a constricted connection with marine water is present which reduces tidal amplitude and therefore mixing. Certainly, the very fresh water immediately under the ice indicates that tidal mixing is insufficient to mix this surface water on a short (though indeterminate) time scale. However, the presence of a basin containing saline water indicates that marine water invaded the lake at some earlier time as also proposed for Transkriptsii Gulf.

Lake 12 contained freshwater, as well as a more saline layer beneath 60 m. It was not determined if this saline water was anoxic, which could indicate a likely depth for the hydraulic connection. Most interestingly, the salinity of the mid-depth brackish water was nearly identical to that over considerable depth ranges in both Transkriptsii Gulf and Lake 14. The close similarity in these salinities could be due to three reasons. Firstly, it could be due to chance. This is unlikely, as it would require that mixing at the marine – freshwater interface in three lakes of quite different size, depth and structure has resulted in water of the same salinity. The second possibility is that all three lakes receive (or received) inputs from the same water mass. This 41 implies that the marine connections for all three lakes coalesce and mix at some point under the ice. Thirdly, it is possible that the connections to Lakes 12 and 14 are not to the marine waters, but rather originate in Transkriptsii Gulf itself. At present, there is not enough information to determine which is the case. Detailed tidal measurements would help determine if this were the case.

The cold water at the base of Lake 12 suggests that saline water at its freezing point reaches the lake (compare this to the increase in temperature with salinity at the base of Transkriptsii Gulf, the brackish basin of Lake Pol’anskogo (Gal’chenko et al., 1995), and Lake 14. The relatively ‘clean’ temperature profile (compared to that of Transkriptsii Gulf) in the region 30-50 m further suggests there is no input in this region. The less saline water at the top of the lake indicates that freshwater balance is currently positive, and that this water is mixing with and diluting the slightly more saline water beneath. The force for this mixing is uncertain, but could possibly be due to an hydraulic connection at these depths.

The profiles of Lake 14 indicate that it is probably similar to Lake Pol’anskogo, in that it has two basins. One, possibly to the western side of the Edisto Moraine where it dips under the surface of the ice, is likely to be fresh, and be the site of the tidal exchange with the ocean, and the second to the east (the basin sampled), is warmer and saline. The shallowness of the increase in temperature and salinity indicates that the ridge separating the two basins is quite close to the surface. It is probable that the water in this basin is marine water that entered the basin during periods of lower freshwater input.

The epishelf lakes of the Bunger Hills are fascinating physical environments, and cry out for further studies of, for example, currents and tidal measurements to clarify mixing processes and the historical messages they can reveal.

4.3 Chemistry

Most of the knowledge of the chemistry of the lakes of the Bunger Hills comes from reasonably detailed surveys conducted by Russian and associated researchers in the late 1980s and early 1990s (Klokov et al., 1990; Kaup et al., 1993; Gal’chenko, 1994). Further data for eight lakes is provided by Barker (1977), and limited data is also available for a small lake near Dobrowolski Station (Markov et al., 1970). Most of this chemistry has dealt purely with the major 42 ion composition. No data are available on, for example, heavy metals concentrations in the lakes or elsewhere.

Klokov et al. (1990) separated the lakes into four categories, summarised in Table 3, depending on their chemistry and history. The lakes that receive direct input of water from the plateau, including Lakes Algae and Dalekoje, had very low concentrations of dissolved ions (electrical conductivity 31-38 and 16-19 µS cm -1 respectively, c.f. White Smoke Lake: 43 µS cm -1). In molar terms (i.e. taking the atomic weight of the elements into account), the dominant cations were + 2+ + 2+ - 2- - Na >Ca >K >Mg , and for anions Cl >SO 4 >HCO 3 . As the general proportions of the ions are approximately those in seawater, the source of the ions is most likely from deposition of marine aerosol on the ice that then enter the melt water, though with the addition of considerably more calcium. Some ionic input would also come from interactions with rock sediments and by dissolution of sub-glacial calcium carbonate deposits.

Table 3. Lake categories based on water chemistry, mixing and catchment characteristics proposed by Klokov et al. (1990).

Category Attributes Examples Lakes with glacial melt Low conductivity, melt water input from Algae Lake, Lake Dalekoje water through flow the glacier. Isolated lakes of glacial Low to moderate conductivity, melt Lake Dolgoe, Lake origin water from land only, but some through Dolinnoje flow. Isolated lakes of sea origin High conductivity, generally closed Lake Polest, Lake Vostochnoye Sea inlet Low salinity, tidal Transkriptsii Gulf, Lake Pol’anskogo

Once isolated from glacial melt water inputs, the salinity of a lake rises due to evaporation. The conductivity of Lakes Dolgoe, Dolinnoje and 17 (Figure 9), was 190 – 1000 µS cm -1, one or two orders of magnitude greater than that of the lakes which still have glacial melt water input. The dominant cations were again in the same order, with the exception that Ca 2+ and Mg 2+ were reversed. This reflects a greater percentage of marine aerosol input and reduced - - calcium carbonate influence. Cl was still the dominant anion, but HCO 3 was more concentrated 2- 2- than SO 4 . Klokov suggested that the relatively low levels of SO 4 may have been due to the 43

precipitation of mirabilite (Na 2SO 4.10H 2O), but these lakes are currently not nearly saline enough for this to happen (in the Vestfold Hills, mirabilite precipitation only occurs in lakes with salinities above about 150 g L -1). Two of these lakes were observed to either have an outlet during the 1999/2000 ANARE visit, or would do so in a normal melt year (Lake Dolinnoje was not visited). If the snow melt input into the lakes had a similar ionic makeup to the polar ice (as would be expected), the higher concentrations must be the result of evaporation during the long residence time of water in the lakes. The large flow through Lakes Algae and Dalekoje have not allowed this to happen to the same extent, but it is interesting to note that Algae Lake, with a much longer residence time and larger surface area (and therefore potential for evaporation) was the more highly mineralised of the two.

To the north of Algae Lake, the lakes were generally far more mineralised (conductivities in the range 1170 – 37990 µS cm -1), with the ionic constitution closing matching that of diluted or concentrated seawater. Klokov et al. (1990) suggested that these lakes had formed as a result of isostatic rebound trapping pockets of seawater as the land rose, a process well documented for the saline lakes of the Vestfold Hills (Burton, 1981). The occurrence of this mechanism in the Bunger Hills is problematic, as many of the saline lakes are positioned above the maximum marine incursion a few metres above current sea level. The salt in these lake must therefore have also come from marine aerosol deposition. There are at least two reasons why these lakes have ended up more saline than those to the south of the Algae Lake: (i) they generally do not have an outlet, and therefore a chance to export salt from the lake basin; and (ii) they are downwind of Rybij Khvost Gulf, which is open water in most summers, and therefore could be an important source of wind blown salt. The saline lakes appeared to be in most cases shallow, with the maximum depth of the largest, Lake Polest, being only 6 m. It is likely, though has to be shown, that none are meromictic.

The final category listed by Klokov et al. (1990) was ‘sea inlets’, which includes Transkriptsii Gulf and Rybij Khvost Gulf. The ionic ratios throughout the water column in Transkriptsii Gulf is virtually identical to that of seawater, suggesting that direct dilution of seawater had occurred. This is consistent with the mixing in the epishelf lakes discussed earlier.

Salinity measurements, recorded to an accuracy of perhaps 5% of the reading with an Otago hand held refractometer, were made for approximately 80 lakes during the 1999/2000 ANARE visit. The general pattern of salinity, shown in Figure 25, was similar to that recorded 44

Southern Bunger Hills N

0-1 g L -1 1-5 g L -1 5-10 g L -1 10-20 g L -1 20-40 g L -1 0 5 -1 km >40 g L

Figure 25: Distribution of surface lake salinities measured during the 1999/2000 ANARE visit. previously, and confirmed the basic division between freshwater lakes to the south of Algae Lake and Transkriptsii Gulf, and more saline lakes to the north. It is clear, however, that some saline lakes occur to the south – the two examples were both in closed basins with little chance of overflow – and that the salinity of the saline lakes was very varied. The salinity will be a function of water flow through the lake and current water level. As mentioned above, water through flow will help to flush salt from a basin (or preclude its build-up in the first place). In closed systems, water level will be approximately inversely proportional to salinity; evidence of low water levels were observed in any lakes in the northern areas, indicating that salinities would currently be high. The maximum salinity recorded was 81 g L -1 in Lake Polest; higher salinities (>130 g L -1) have been noted in lakes in the northern section of the Bunger Hills by previous workers (Melles, 1994). These maximum salinities are still well beneath those recorded for some lakes in the Vestfold Hills (Burton, 1981; Gibson, 1999).

The pH of the lakes was measured to be between 6.9 and 8.8 (Klokov et al., 1990), which are unexceptional values. The most interesting pH was recorded at the interface between anoxic and oxic water in Transkriptsii Gulf, which was the lowest in the lake. This was also not unexpected; the acidity stems from the oxidation of sulphide at the top of the anoxic zone.

45

Nutrient (nitrate and phosphate) concentrations in the lakes were very low (5 – 10 µg L -1), indicating oligotrophic conditions (Klokov et al., 1990: note that the units for nutrient concentrations given in this paper are incorrect – they should µg L -1 rather than mg L -1 (E. Kaup, personal communication)). Total phosphorus concentrations were also low (circa 10 µg L -1). These concentrations are similar to those recorded in lakes of the Vestfold Hills (Perriss and Laybourn-Parry, 1997; Rankin et al., 1999). Far higher concentrations of total phosphorus (>800 µg L -1) were recorded in the anoxic water of Transkriptsii Gulf resulting from remineralisation of sedimented organic material. Silica was present in concentrations between 10-30 mg Si L -1, reflecting some leaching of the rock basins in which the lakes occur. Similar concentrations have been recorded in the lakes of the Vestfold Hills (Rankin et al., 1999).

Klokov et al. (1990) reported that dissolved oxygen was near saturation in lakes that were fully or partially ice-free, though under ice cover some supersaturation (maximum: 125%) was observed. Similar supersaturation is well known in lakes of the Vestfold Hills and elsewhere in Antarctic (Wharton et al., 1986; Rankin et al., 1999). Anaerobic water has been recorded at the base of Transkriptsii Gulf and the saline basin of Lake Pol’anskogo (Klokov et al., 1990; Gal’chenko et al., 1995). Two oxygen profiles were recorded during the 1999/2000 ANARE visit in Lake Polest and Transkriptsii Gulf (to a depth of 30 m). The profiles, shown in Figure 26, indicate supersaturation of up to 130% in Transkriptsii Gulf, and 220% in Lake Polest. No anoxic water was present in Lake Polest, consistent with absence of meromixis indicated by the salinity and temperature profiles. In both environments, photosynthesis would have resulted in production of oxygen, which could not equilibrate with the atmosphere due to the ice cover present. The presence of undersaturated waters below about 25 m in Transkriptsii Gulf indicates that remineralisation of organic matter produced in the surface waters outweighs primary production at these depths. The structure apparent in the oxygen profile match up with stratification indicated by temperature changes, showing that significant mixing is not occurring in these surface waters.

4.4 Biology

The biology of the lakes of the Bunger Hills has been studied only cursorily. The information available is scattered, and the lists of species present, for example, should be treated very carefully as a result of difficulties with identifications and changes in taxonomy and nomenclature. The following sections discuss what is known about organisms that live in the lakes, ranging in size and complexity from bacteria to copepods. 46

(a) (b) Dissolved oxygen saturation (%) Dissolved oxygen saturation (%)

90 100 110 120 130 160 180 200 220 240 0 0 Ice Ice 5 1

10 2

15 3 20 Depth Depth (m) 4 25 5 30 6 13 14 15 16 17 18 19 20 12 13 14 15 16 17 18 19 20

Dissolved oxygen (mg L -1 ) Dissolved oxygen (mg L -1 )

Figure 26: Profiles of dissolved oxygen recorded in (a) Transkriptsii Gulf (to a depth of 30 m) and (b) Lake Polest.

4.4.1 Non-photosynthetic bacteria

Few studies of the bacteria (not including cyanobacteria, which are discussed in Section 4.4.2) of the lakes of the Bunger Hills have been made. None of these studies has been taxonomic in nature, and thus there is no knowledge, except in the broadest sense, of the species of bacteria which inhabit the lakes. However, a number of important bacterial processes have been found to occur in water column and/or sediments of both permanently and perennially ice covered lakes, including sulphate reduction, methanogenesis and methane oxidation (Gal’chenko, 1994), which imply the presence of sulfate-reducing, methanogenic and methylotrophic bacteria. All these bacteria occur in other lakes in Antarctica and throughout the world. Even though methane production was detected, the rate of oxidation was such that methane did not accumulate in the water column. A series of papers has also appeared that use samples collected in the Bunger Hills to develop or test new biochemical methods (e.g. the detection of the ribulose bisphosphate carboxylase gene (Chernykh, 1995), and the detection of methylotrophs (Beliaev et al., 1995)).

4.4.2 Photosynthetic bacteria and algae

The water column of most of the lakes was observed during the 1999/2000 ANARE visit to be very clear, implying low concentrations of chlorophyll a and therefore photosynthetic organisms within the water column. Klokov et al. (1990) reported chlorophyll a concentrations of 47 less than 1 µg L -1 in nearly all the lakes they studied, supporting this observation. Given the ice- free nature of the lakes and clarity of the water, this suggests very low nutrient concentrations, confirmed by the data given above. The only lake with a higher concentration of chlorophyll a amongst those studied by Klokov et al. (1990) was Lake Dalekoje, which possibly received nutrients from its input of glacial flour. The concentration of chlorophyll a was similar in the saline lakes of the region (Kaup et al., 1993), being generally less than 1 µg L -1 except for values up to about 1.8 µg L -1 on occasions in Lakes Vostochnoye and Zapadnoye. The paucity of the flora of the water column was supported by Hay (in Ledingham, 1986a), who went so far as to suggest that phytoplankton were totally lacking. The generally low chlorophyll a concentrations in the lakes of the Bunger Hills were confirmed in samples collected during the 1999/2000 ANARE visit from Lake Polest and Transkriptsii Gulf. The concentration range in the former was 0.15 – 0.54 µg L -1, and in the Gulf (from depths between 5 and 50 m) 0.36 – 0.57 µg L -1. The few neritic phytoplankton species identified by Kaup et al. (1993) in a study of the saline lakes of the Bunger Hills, listed in Appendix 2, were all diatoms that were also common in the nearby marine environment.

Primary productivity measured by incorporation of radio-labelled carbonate into organic material was measured by Kaup et al. (1993) in the water column of a number of lakes, including Lakes Polest, Vostochnoye and Zapadnoye. Productivity in these saline lakes, which ranged from 13-171 mg C m -3 d -1, was generally an order of magnitude higher than in freshwater lakes of the Bunger Hills (0.6-19 mg C m -3 d -1). These rates compare to circa 20 mg C m -3 day -1 in Ace Lake in the Vestfold Hills (Rankin et al., 1999; Swadling and Gibson, 2000). Highest productivity was observed in the warm hypolimnion of Lake Polest during a period of seasonal stratification. Evidence of photoinhibition of productivity in the highly illuminated surface waters was apparent, as has been observed elsewhere in Antarctic lakes. Marked interannual variations in productivity occurred in Lake Vostochnoye, probably related to the presence or absence of ice and its effect on the underwater light field. Similar interannual variability is commonly observed in Antarctic lakes. A slightly wider range of production rates (0.08 – 326 mg C m -3 day -1) was given by Gal’chenko et al. (1995), who also noted a general relationship between productivity and water salinity. The highest value was again recorded in the summer stratified water of Lake Polest.

In contrast to the water column, the benthic flora of the lakes is abundant. Nearly all lakes have thick microbial mats consisting of a community of species including cyanobacteria, chlorophytes, diatoms, mosses, and a suite of heterotrophs which graze upon this material (Figure 27). The 48 appearance of these mats was dependent on a number of factors, including depth and water salinity. Mats were usually less developed close to the margins of the lake where ice formation would provide a physical hindrance (both in terms of temperature and abrasion) to mat development. In deeper water much thicker mats (up to 5 cm thick) were observed. In summer, portions of these mats (‘lift-off mats’) are often blown to the downwind end of the lake, where they accumulate. In general, the surface few millimetres of the mats are either orange or brown in colour, beneath which they are green. These colours are due to the presence of different species of cyanobacteria, some of which produce orange UV-screening compounds and are adapted to living in exposed situations, and others which lack these pigments, are green, and take advantage of protection from UV afforded by the pigmented organisms. Lower salinity lakes generally have more loosely aggregated mats with orange surfaces, but as salinity increases, the mats tend to be browner and more cohesive in texture. Microbial mats are well known from lakes throughout the Antarctic (Vincent, 1988) and the Arctic (Vincent et al., 2000), and in many cases the mats are the site of the majority of the primary productivity in the lakes (Heath, 1988).

Little is known about the biological make up of the microbial mats in the lakes of the Bunger Hills. Korotkevich (1964c) and Vialov and Sdobnikova (1961) reported a number of species of cyanobacteria and diatoms, as did Barker (1977), who also listed chlorophytes (see

Figure 27: Brown microbial mat between rocks close to the margin of a saline lake. January 2000. A small piece of detached mat can be seen to the left of the large rocks at the bottom left and in the centre of the Figure. 49

Appendix 2 for a list of species recorded in the Hills). In particular, there is little knowledge of the effect of salinity in structuring these communities. Of some interest is the occurrence of desmids in a number of lakes; desmids are generally scarce in Antarctic lakes (Ellis-Evans et al, 1998).

4.4.3 Fauna

The lake fauna of the Bunger Hills is of considerable importance, as the area is the type location for at least five species (see Appendix 3 for a list of species for which the type location is in the Bunger Hills). A number of these species, especially three species of copepod, are of particular biogeographic interest. This section describes the little that is known about the fauna of the lakes. Note: Heterotrophic protists, tardigrades and nematodes, which occur in both lacustrine and terrestrial (e.g. moss) environments, are discussed under the heading Terrestrial Fauna (Section 6.1-3).

4.4.3.1 Rotifers

Korotkevich (1964c) listed seven species of rotifer in samples collected from Bunger Hills lakes (see Appendix 4 for a list), to which Kutikova (1964b) added another species. Rotifers were also observed in samples collected during the 1977 ANARE visit, but Barker (1977) gave no identifications. Sudzuki (1979) recorded the presence of rotifers in preserved samples collected by during the 1977 ANARE visit, but could not identify them to species.

The rotifer Notholca verae was described from material collected from Algae Lake (Kutikova, 1964a), and has subsequently been reported from other lakes in this region (Korotkevich, 1964c) as well as the Vestfold Hills (Everitt, 1981). Sudzuki (1988) made a careful study of Antarctic members of the genus Notholca , and concluded that Notholca verae was a good species. More recent studies by Dartnall (personal communication) indicated that the material from the Vestfold Hills was not identical to that described by Kutikova (1964a), and is possibly a separate species. It is probable that Notholca verae and the related species in the Vestfold Hills are endemic Antarctic species that have survived on the continent through periods of more intense glaciations, possibly in refugia such as epishelf or supraglacial lakes.

50

Microbial mat samples collected during the 1999/2000 ANARE visit included the following species: Lepadella patella , Notholca sp., Adineta grandis , Adineta sp., Habrotrocha sp., Philodina gregaria , and Philodina sp. (H. Dartnall, personal communication). Of these, at least four are new to the Bunger Hills (Appendix 4).

4.4.3.2 Platyhelminthes

Hay (in Ledingham, 1986a) searched for platyhelminthes (flatworms) in the lakes of the Bunger Hills during the 1986 ANARE visit. Cocoons were found in a small tarn to the east of Edgeworth David camp, but no adults were observed in the benthic community of any of the many lakes visited. No further information about platyhelminthes in the lakes of the Bunger Hills is available.

4.4.3.3 Copepods

Two species of copepod have been described from the lakes of the Bunger Hills, both of which are of great biogeographical interest. The cyclopoid Acanthocyclops mirnyi was described from samples collected from Algae Lake on an early Russian visit (Borutzky and Vinogradov, 1957). This species also occurs in numerous lakes in the Vestfold Hills (Laybourn-Parry et al., 1995; Dartnall, 2000; K. Swadling, unpublished data), where it is restricted to lakes of low salinity (<4 g kg -1), and the Larsemann Hills (Dartnall, 1995). Other species of the genus Acanthocyclops have been recorded from islands in the Southern Indian Ocean, Macquarie Island and Australia, but as yet there has been no critical examination of affinities between these species, and therefore possible routes of ancient or post-glacial colonisation. Little is known of the distribution or life history of Acanthocyclops mirnyi ; an adult female bearing eggs captured in March was pictured in Barker (1977), suggesting that the species wintered at an early stage (possibly even as diapause eggs) before developing to adults in summer. Korotkevic (1964c) reported capturing only juvenile stages in Algae Lake in January 1957, though no further details were given. In the Vestfold Hills, ovigerous females were present in late February (D. Grace, unpublished notes). It has been suggested that the Vestfold Hills population is essentially benthic (Laybourn-Parry et al., 1995), but a recent study of this species shows morphological characteristics typical of neritic species (J. Reid, personal communication).

51

The second species present in the lakes of the Bunger Hills is the calanoid Gladioferens antarcticus (Bayly, 1994). This species has been reported previously only from White Smoke Lake, where adults were present in March. This observation again suggests a wintering strategy based on an early stage. The copepods were found to be most common near the base of the lake at 90 m. The water from the lake is particularly clear, with very low chlorophyll levels. Some microbial mat was recovered during the 1999/2000 ANARE visit, suggesting that this may be the food source for the animals.

The biogeography of Gladioferens antarcticus is of particular interest (Bayly, 1994). The genus Gladioferens is euryhaline (i.e. capable of surviving over a wide salinity range) but not marine, and, apart from the Bunger Hills species, is distributed along the southern coast of Australia and New Zealand. The genus has not been recorded from the Southern Ocean. How the copepod reached Antarctica is unknown, but Bayly (1994) suggested that it could be a relict species from prior to the break up of the Australia-Antarctica super-continent. The species may have been able to survive in epiglacial lakes like White Smoke Lake (Bayly and Burton, 1993; Bayly, 1994), which might be present throughout glacial periods. Subsequent dispersion would be difficult, limiting species to relatively small areas.

Few records of copepods in the lakes of the Bunger Hills have been published apart from the two descriptions. Korotkevich (1964c) recorded Acanthocyclops mirnyi in one other deep, freshwater, but unidentified, lake. Barker (1977) found Acanthocyclops mirnyi in the outflow of Algae Lake, and Hay (in Ledingham, 1986a) reported abundant but unidentified copepods in samples from lakes south of Algae Lake, especially in the series of lakes that lie in a prominent fault line that parallels Lake Dolgoe to the south. Klokov et al. (1990) reported the existence of an orange species circa 1 mm in length, which they assumed (correctly) to be Acanthocyclops mirnyi , in many of the freshwater lakes. Finally, Bayly (1994) reported the occurrence of Acanthocyclops mirnyi in samples from White Smoke Lake from which he described Gladioferens antarcticus .

Samples containing at least two species of copepod were obtained from 11 lakes during the 1999/2000 ANARE visit (see Figure 28 for the sampling locations). Copepods were particularly abundant in Transkriptsii Gulf and Lakes 12 and 14, but scarce in the low chlorophyll a lakes along the southern margin of the Bunger Hills (e.g. Lake Pol’anskogo, White Smoke Lake) and in the large freshwater lakes (Lakes Algae and Dolgoe). In Transkriptsii Gulf, 52

Figure 28: Locations from which copepods were collected in 2000. All sites had populations of Acanthocyclops mirnyi , and those with open boxes also had Gladioferens antarcticus . maximum numbers appeared to be associated with the oxic-anoxic interface near 88 m, where both juveniles and adult stages (females with eggs) were present. The low numbers caught in some of the other lakes might reflect the fact that both species appear to prefer living near the deepest part of the lake (Korotkevich, 1964c; Bayly, 1994), which may not have been sampled.

Acanthocyclops mirnyi was by far the dominant species in most of the lakes, with only a few individuals of Gladioferens antarcticus recorded from Pol’anskogo and White Smoke Lakes, Lake 12 and Transkriptsii Gulf. One interesting aspect is that individuals of Acanthocyclops mirnyi from the Bunger Hills were orange in colour, whereas those of the Vestfold Hills are colourless.

A further species of calanoid copepod, Paralabidocera separablis , has been described from the marine ecosystem of the Bunger Hills (Brodsky and Zvereva, 1976). This species has not been recorded elsewhere, but is related to a common and widespread inshore species, Paralabidocera antarctica , which also occurs in saline lakes of the Vestfold Hills. It is unlikely that this species was collected during the current visit, as the marine environment was not sampled. If Paralabidocera separablis is endemic to the marine ecosystem of the Bunger Hills, it is also of particular biogeographic importance, and genetic studies could provide insights into the 53 age of the isolated Bunger Hills/Shackleton ice shelf ecosystem. An ascidian, Cnemidocarpa zenkevitchii, has also been described from these marine waters (Vinogradova, 1958).

5. Terrestrial flora

The terrestrial flora of the Bunger Hills is limited to algae, fungi, lichens and mosses, with the possibility that a liverwort also occurs. Studies of the flora have been limited to development of species lists and cursory distribution maps. At least two new species (the moss Bryum korotkevicziae (Savich-Lyubitskaya and Smirnova, 1964), which has since been reduced to synonymy with Bryum pseudotriquetrum (Seppelt and Kanda, 1986)) and a fungus, Dactylospora dobrowolskii (Olech and Alstrup, 1996)), and one new form ( Grimmia doniana f. antarctica (Kuc, 1969)) have been described from the Bunger Hills.

5.1 Terrestrial algae

Algae grow in many terrestrial environments, though all of course require water. The sections below deal with observations of a number of these environments.

5.1.1 Stream algae

Most of the major stream beds in the southern Bunger Hills were heavily colonised by thin, black cyanobacterial crusts probably containing species of the genera Nostoc and Schizothrix . Barker (1977) recorded six species of cyanobacteria, a desmid and seven species of diatoms in samples collected from the stream connecting Algae Lake and Transkriptsii Gulf (Appendix 2). Many of the obvious streams in the southern Bunger Hills were not flowing during the 1999/2000 ANARE visit (see comments above about the intensity of the melt during this period), but may have done so later in the summer, thus providing moisture for the algae. It is probable that greater stream flow has occurred in previous years of greater melt. Black cyanobacterial crusts also occurred extensively around the margins of most, if not all, lakes (for example, see Figure 16), reflecting the relatively damp microclimate.

Occasional patches of green algae were seen in streams during the 1999/2000 ANARE visit, and one sample was collected from near White Smoke Lake. Prasiola crispa , a species common in high nutrient conditions elsewhere in Eastern Antarctica, was not observed. This 54 absence is attributable to the lack of Adélie penguin rookeries in the area which are typically the source of the required nutrients.

5.1.2 Epilithic algae

Epilthic algae were observed to be widespread throughout the southern Bunger Hills, but no attempt was made to map these occurrences, or to collect any samples for identification. Epilthic algae were evident as black stains on rock faces, especially in areas of summer seepage. Such conditions were prevalent to the south of Algae Lake, where more snow appeared to accumulate and melt, and the relief was more rugged.

5.1.3 Sublithic algae

No systematic search for sublithic algae was made during the 1999/2000 ANARE visit. Random overturning of light coloured rocks (generally small quartz cobbles), however, revealed rich sublithic communities (Figure 29).

Figure 29: A quartz rock showing abundant and stratified sublithic algae.

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5.1.4 Soil algae

Barker (1977) reported that two species of green algae were isolated from his site 3 (see Figure 4). One isolate was identified as belonging to the genus Trebouxia .

5.2 Fungi

Limited studies of the fungi of the Bunger Hills have been reported on by Barker (1977) and Olech and Alstrup (1996). Two soil samples collected from Barker’s site 3 (see Figure 4) contained nine identifiable species, along with seven isolates which could not be determined. The later report dealt with a number of lichenocolous (i.e. living on lichen) species, including the description of Dactylospora dobrowolskii . The species identified in these studies are listed in Appendix 5. Olech and Alstrup (1996) considered it likely that many further lichenocolous species would be discovered in their samples on further studies, and also highlighted the difficulties in separating lichens (consortia of fungi and algae) from free-living fungi.

5.3 Lichens

The lichen flora of the Bunger Hills has received relatively little study. Early Russian reports noted ‘black, white and grey species’ (Markov et al., 1970), and a series of reports in Russian by Golubkova and Savich covering the lichens of East Antarctica probably contained information about samples collected in the Bunger Hills, though at present only one of these, dealing with the Acarosporaceae, can be confirmed (Golubkova and Savich, 1965). Barker (1977) recorded 17 species (3 only to genus) of which 13 have also been recorded in the Vestfold Hills (see Appendix 6 for a list of species recorded in the Bunger Hills). Many of the species have also been recorded in the Windmill Islands, the next major ice-free area to the east of the Bunger Hills and an area renowned for its abundant lichen and moss flora. The next study of the lichens of the Bunger Hills was that of Olech (1989), who gave a species list, a distribution map, and short ecological comments. Thirty-three species were included in this list, including many species in common with the list of Barker (1977), the flora of the Vestfold Hills and the flora of the Windmill Islands (Appendix 6). A further four species were listed in a supplementary paper (Olech and Alstrup, 1996). A Russian report of lichen studies in the Bunger Hills has appeared (Andreev, 1991), containing a list of at least 41 species from 22 genera and nine families. It was noted that 13 of the species were endemic to Antarctica. Only an abstract of this paper was seen 56 during the preparation of this report, and the list of species recorded was not available. Occasional mentions of lichens have been made in other, often geomorphological, papers, the most interesting of which was a study of the thallus diameter of one of the most widespread species in the Bunger Hills, Buellia frigida (Bolshiyanov et al., 1991). This paper unwittingly gives a good distribution map for the species in the southern Bunger Hills.

Some of the lichens recorded in the Bunger Hills been reported only from distinctly different biogeographical zones. Two species listed by Olech (1989) were noted to be new to the flora of continental Antarctica: Arthonia subantarctica , originally described from Bouvet Island, and Carbonea vorticosa were found at 3 sites and one site respectively. Physcia dubia , a species previously recorded from temperate to boreal regions of the northern hemisphere (but also with an anomalous record from Lützow-Holmbukta, Antarctica: see comments in Seppelt, 1986b), was collected from five sites (this species was also collected in the Bunger Hills by Adamson in 1986 (R. Seppelt, personal communication)). Olech and Alstrup (1996) listed another three species new to the Antarctic continent: Buellia pulverulenta , Caloplaca saxicola (previously recorded in the South Shetland Islands), and Leprocaulon subalbicans (also previously recorded in the South Shetland Islands). If these identifications are correct, the Bunger Hills contain a series of unusual species for a continental Antarctic site. However, with the difficulty in accurately identifying lichen species as well as the fluid state of Antarctic lichen taxonomy, these identifications must be treated with caution until an independent specialist can look at samples.

During the 1999/2000 ANARE visit, lichens were found to be common over much of the southern Bunger Hills. The presence or absence of lichens was noted in 1 km 2 quadrats (based on the UTM coordinate system; see Figure 30 for a map of the quadrats visited). An attempt was made to visit as many quadrats as possible. Figure 31 shows those quadrats surveyed in which lichens were present, as well as sites from which lichens have been recorded in previous studies (Barker, 1977; Olech, 1989; Bolshiyanov et al., 1991; Olech and Alstrup 1996). It should be noted that the absence of lichens in any particular quadrat is not absolute, as no quadrat was surveyed thoroughly. It is therefore possible that lichens were missed in some of the quadrats for which lichens are shown to be absent. However, the indication of the presence of lichens can be relied upon. Except for areas to the north and east of Edgeworth David Base, nearly all quadrats had at least some lichens. The absence of lichens to the north was probably related to increased soil salinity, rock weathering (D. Gore, personal communication), or exposure to saline

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Figure 30: UTM-based kilometre quadrats surveyed during the 1999/2000 visit.

Figure 31: Quadrats in which lichens were observed. Also noted are the positions of previous reports of lichens from Barker (1977: triangles), Olech (1989: small circles), Bolshiyanov et al. (1991: squares) and Olech and Alstrup (1996: small circles). aerosols from not only the marine waters of Rybij Khvost Gulf but also the saline lakes of the area, much in the same way that the distribution of lichens in the Vestfold Hills is strongly influenced by these factors. Available water could also be an important controlling factor, as there 58 appeared to be less snow build up in the northern part of the southern Bunger Hills compared to areas closer to the Apfel Glacier to the south, and therefore less available melt.

In the north, lichens were limited to scattered colonies of Buellia frigida growing on boulders often with a westerly aspect (i.e. protected from the prevailing easterly winds). South of a line passing through Edgeworth David Base and a few hundred metres to the north of Izvilistaja Bay and Algae Lake, abundance and diversity increased. To the north, communities containing species other than Buellia frigida occurred in melt streams emanating from snow banks, but further south they became more widespread. The richest sites were generally found in low-water flow stream beds, especially associated with periglacially-formed cracks, and on south facing slopes relatively close to the southern margin of the Bunger Hills, where coverage in some areas reached 25 - 50%. Lichens were also widespread on a number of isolated nunataks to the south of the main Bunger Hills. Interestingly, some stretches of the southern margin appeared to be lichen- free for hundreds of metres, suggesting very recent deglaciation.

Lichen diversity also increased from north to south. Around Edgeworth David Base, only one species ( Buellia frigida ) was reasonably common, whereas to the south of the line mentioned above the number of species increased. Maximum number of species were found in a relatively narrow band (maybe 1-2 km wide) at the southern margin of the Hills (Figure 32). It is difficult at this stage to estimate the number of species present at some of the richer sites, but it was probably of the order of 20.

An attempt was made to map the distributions of a number of readily recognisable species or genera. It should be recognised that none of the people undertaking this survey were lichen specialists, and maps given below should be considered a starting point for any subsequent investigations, and should not be considered either accurate or complete.

Buellia frigida

This crustose species is quite readily recognised, especially in the those areas where it is one of the few species present. It was characterised by an effigurate lobed margin, grey to blackish grey in colour, with a lighter zone a few millimetres in from the edge of the thallus. Inside this zone the colour was again darker. In older colonies, the centre of the thallus had often

59

Figure 32. A site showing a wide diversity of lichens and mosses located close to the southern edge of the Bunger Hills.

Figure 33: Distribution of Buellia frigida recorded during the 1999/2000 ANARE and previous visits. See Figure 31 for an explanation of the symbols used.

60 been lost by abrasion. This species was the most widespread of those surveyed in this study (Figure 33, which also shows localities for this species recorded by Barker (1977), Olech (1990), and Bolshiyanov et al. (1991)), being present in nearly every quadrat lichens were found. The Bunger Hills are therefore similar to the Vestfold Hills, where Buellia frigida is the most widely occurring species (Seppelt, 1986b). In general, B. frigida was found on medium to large boulders, and was the dominant species in the immediate vicinity of the Edgeworth David camp.

Physcia caesia

The grey foliose species Physcia caesia (syn. Parmelia coreyi ) occurred over most of the southern part of the area surveyed (Figure 34). It was most abundant in melt streams, where it formed a loose association with Umbilicaria aprina, Rhizocarpon geographicum and Pseudephebe minuscula. Physcia caesia was also common in periglacial crack systems, again in association with same three species. The apparent gap in the distribution in the centre of the area surveyed is probably an artifact.

Figure 34: Distribution of Physcia caesia recorded during the 1999/2000 ANARE and previous visits. See Figure 31 for an explanation of the symbols used.

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Pleopsidium chlorophanum

This bright citron yellow (and therefore readily identifiable) crustose species was found mainly along the southern ice margin growing in species-rich communities with good water supply (Figure 35). It was also found between the southern shore of Lake Algae and Lake Pt’ich’je. This species was the most sporadically distributed amongst those surveyed, and appeared to have the greatest need for the ‘right’ conditions – large, frost-shattered rocks providing protection and a good source of water from snow banks or ice from above. This species is rare in the Vestfold Hills, but locally common in the Bunger Hills.

Pseudephebe minuscula

This black, minutely fruiticose species was quite easily identified. It was abundant throughout the southern part of the area surveyed, but was observed at only one or two locations to the north of Algae Lake (Figure 36). In general, Pseudephebe minuscula grew in association with Umbilicaria spp. and Physcia caesia in drainage channels in periglacial landforms. In transects towards drier and less lichen-rich areas, Pseudephebe minuscula dropped out after

Figure 35: Distribution of Pleopsidium chlorophanum recorded during the 1999/2000 ANARE and previous visits. See Figure 31 for an explanation of the symbols used. 62

Figure 36: Distribution of Pseudophebe minuscula recorded during the 1999/2000 ANARE and previous visits. See Figure 31 for an explanation of the symbols used.

Physcia caesia , but before Umbilicaria aprina . Towards the south it was more likely to be seen growing on rock slabs away from such environments. Pseudephebe minuscula is a rare species in the Vestfold Hills (Seppelt, 1986b), but is common and widespread in the southern Bunger Hills.

Rhizocarpon geographicum

Rhizocarpon geographicum is an easily identifiable crustose species with a yellow areolate thallus with black margins and black between the areoles. In the Bunger Hills it is relatively cryptic, as it often occurred in small colonies in nooks and crannies in rocky areas. Rhizocarpon geographicum was, however, found to be widely distributed throughout the southern portion of the study area, often growing in association with Umbilicaria aprina , Pseudephebe minuscula and Physcia caesia (Figure 37).

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Figure 37: Distribution of Rhizocarpon flavum recorded during the 1999/2000 ANARE and previous visits. See Figure 31 for an explanation of the symbols used.

Umbilicaria spp.

At least two species of the foliose genus Umbilicaria are present in the Bunger Hills: Umbilicaria aprina and Umbilicaria decussata . These species are relatively easily separated in the field, but are mapped together in the distribution map. Umbilicaria aprina (with the undersurface of the thalli having hair like structures or rhizines) was the second-most widespread species found in the survey (Figure 38). It occurred widely in melt streams under snow banks, as well as very abundantly in periglacial cracks (see Figure 39). In many areas, the cracks on all sides of a polygon in areas of patterned ground would be heavily colonised by this species, while the centre of the polygon would be lichen-free. Individual thalli were in most cases relatively small (up to 2 cm across), but occasionally, especially in well watered areas, reached 10 cm or more. Umbilicaria decussata (which lacks undersurface rhizines) was observed most commonly in areas close to the rock-ice boundary on the southern margin of the Bunger Hills. A possible third species, with relatively small, dark (nearly black) thalli with hairs on the under surface, was also present in this region. In some areas close to the ice-rock margin Umbilicaria was very common, reaching up to 25-50 % coverage of slabs on southern-facing rock steep hillsides above, for example, Lakes Pt’ich’je and Pol’anskogo 64

Figure 38: Distribution of Umbilicaria spp. recorded during the 1999/2000 ANARE and previous visits. See Figure 31 for an explanation of the symbols used.

Figure 39: Periglacial crack with melt stream inhabited by a rich lichen and moss flora including abundant Umbilicaria .

.

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Usnea spp.

Usnea spp. (probably including Usnea antarctica and Usnea sphacelata ), which were easily identified by their fruiticose growth habit, were sparsely distributed along the southern margin of the southern Bunger Hills (see Figure 40). Colonies were often well hidden in protected spots, but most consisted of numerous, closely-packed thalli (Figure 41). In general, Usnea was not found to colonise the beds of melt streams, but rather occurred on rock slabs in areas that would receive some melt from summer melt water from local snow banks. Usnea was abundant on a number of small nunataks located to the south of the main rock area. The records of Usnea sp. at Barker’s site 12 and other sites well away from the southern margin of the Bunger Hills need to be confirmed. At present, these localities appears to be well away from the area in which this species is most common.

In summary, the general distribution of lichens derived from this partial survey, as well as previous surveys, suggests that while some species are capable of surviving over a (relatively) wide range of environmental conditions ( Umbilicaria aprina , and especially Buellia frigida ), others are restricted to much more limited conditions ( Usnea spp., Pleopsidium chlorophanum ). The occurrence of these more ‘sensitive’ species close to the southern Bunger Hills-Apfel Glacier

Figure 40: Distribution of Usnea spp. recorded during the 1999/2000 ANARE and previous visits. See Figure 31 for an explanation of the symbols used. 66

Figure 41: Usnea spp. growing with other lichens in a protected site close to the southern Bunger Hills/Apfel Glacier margin. interface, as well as the apparent maximal diversity in this region, suggests that the most preferable conditions (in terms of weather conditions, water and nutrient availability, and substrate) for lichen growth occurs in this area. However, it must be reiterated that a survey by a trained lichenologist is necessary to confirm the general distributions given above as well as the conclusions drawn.

Two ecosystems appeared particularly important for lichen growth: melt streams, which were often associated with periglacial cracking; and south-facing rock slopes close to the Apfel Glacier. The first were often very heavily colonised by a selection of species, though the intensity of the colonisation generally decreased towards the north. Most of the cracks held liquid water when visited, even if they were not part of an active stream system. North of Algae Lake little periglacial activity was apparent, and, apart from Buellia frigida , lichens were restricted to melt streams emanating from larger snow banks. The south-facing rock slopes close to the Apfel glacier were typically quite steep, and consisted of large, frost-fractured, often unstable rocks. Both exposed surfaces and protected interstices harboured abundant lichens and a wide species diversity.

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5.4 Mosses

Mosses are also widely distributed throughout the southern Bunger Hills, and have been the subject of a number of previous studies. Early work included the comments of the occurrence of Sarconeurum glaciale , Grimmia plagiopodia , Pohlia nutans and Bryum algens in the Bunger Hills in a wider, east Antarctic context (Savich-Lyubitskaya and Smirnova, 1966, 1971a, 1971b, 1972), and the description of Bryum korotkevicziae in samples dredged from a depth of 36-39 m in Algae Lake (Savich-Lyubitskaya and Smirnova, 1964). This aquatic species was characterised by long stems and sparsely distributed leaflets, but more recent studies have shown that it is merely an aquatic growth form of Bryum pseudotriquetrum , and thus should not be considered a separate taxon (Seppelt, 1986a). Kuc (1969) recorded four species in samples collected in 1959, including some rarely reported elsewhere in Antarctica. Amongst these species was the description of a new form: Grimmia doniana f. antarctica . Barker (1977) listed 3 species of moss from samples collected during the 1977 ANARE visit, as well as noting a further, unidentified species. Olech (1989) provided limited records for two species. A list of all species recorded in previous studies of the southern Bunger Hills is given in Appendix 7. This list should be treated with caution, however, until confirmation by independent bryologists is made due to difficulties with taxonomy of the Antarctic bryoflora. Finally, unidentified mosses have also been recorded from Geographer’s Island (Melles, 1994) in the northern Bunger Hills, and noted in sediment cores taken from lakes, where they are an important indicator of lacustrine, as distinct from marine, conditions (Bolshiyanov et al., 1991; Verkulich and Melles, 1992).

During the 1999/2000 ANARE visit, moss was found to occur widely throughout the area visited. Figure 42 shows a distribution map of all moss species on the same kilometre grid used for mapping the lichens. The comments regarding this map are the same as for the maps of lichen distributions as above. No attempt was made to determine the species of moss present.

As for the lichens, the distribution of mosses appeared to be limited by the presence of salt in the region to the north and east of Edgeworth David Station (Vertoletnyj and Krylatyj Peninsulas), as well as to the north of Algae Lake near Dobrowolski Station. Lack of melt water availability in much of this region could also be a factor, as there appeared to be less accumulation of wind-blown snow. In general, mosses were common and appeared in better condition in the south of the study area; the moss towards the north was often moribund, heavily

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Figure 42: Quadrats in which mosses were observed. Also noted are the positions of previous reports of mosses from Barker (1977: triangles) and Olech (1989: circles), and the location of the two lakes from which samples of aquatic moss were recovered (stars). lichenised (often by a yellow lichen), or had a cyanobacterial crust covering it. The most verdant mosses were found in melt streams emanating from snow banks, or in streams between lakes (Figure 39). Of particular importance were drainage systems that included periglacially-formed cracks, which often provided water as well as physical protection. Such cracks were widespread in the southern section of the area studied, but became rarer to the north. Some continuous moss beds up to 1 m 2 in area were observed in areas, but in no place were the larger areas prominent in the Windmill Islands present. In strongly flowing streams (either now or sometime in the past), the mosses were restricted to the margins of the stream where flow was reduced (this is discussed further below). Mosses were also present in moraines along the margins of the ice-free land, noticeably in areas were lichens were sparse or absent.

Samples of aquatic mosses were recovered from two lakes (Figure 42). In the sample from Lake Dolgoe, the individual stems were dispersed through thick, gelatinous cyanobacterial mat, while at the other lake, a bundle of stems interspersed with small amount of mat material was found washed up on the shore. Collections were made from both sites. In both cases, the moss had clearly grown at depth, but had been removed from its growth position, possibly by 69 buoyancy associated with production of oxygen by the associated cyanobacteria as well as the moss itself.

5.5 Liverworts

No liverworts were recorded during the 1999/2000 ANARE visit to the Bunger Hills, nor have they been recorded previously. However, one species ( Cephaloziella varians ) has been found in the Windmill Islands and the Larsemann Hills. Considering the richness of the flora in the Bunger Hills, it is possible that this species occurs there as well.

5.6 Spatial zonation of flora in stream beds and around lakes

Consistent trends were observed in the distribution of lichens, mosses and cyanobacterial crusts in stream beds and around lakes throughout the Bunger Hills. A cross section of a typical stream bed is shown in Figure 43a. Lichens dominated the vegetation away from the thalweg in areas that would receive water from spray, but were not submerged. Closer to the centre of the stream mosses were prevalent, but in the centre of the stream, where flow would be consistently strongest, mosses and lichens were absent, and black cyanobacterial crusts were the only apparent vegetation. This distribution clearly reflects the habitat requirements of the different vegetation types. Lichens have less of a requirement for water than mosses, and are possibly more prone to damage by strong flow. Therefore they are concentrated at the margins of the streams. Mosses require greater water availability, but appear to be unable to withstand the stronger water flows in the thalweg. Cyanobacterial crusts can survive in areas of low water abundance (e.g. in an outer ring around lakes, see below), and thus there is no requirement to be in the area of greatest water flow in the middle of the stream. It is more likely that these crusts are more able to withstand the physical forces experienced in this zone.

Two vegetation zones occurred around the lakes of all salinities (Figure 43b). An outer ring of black cyanobacterial crust (presumably containing Nostoc and Schizothrix ) encircled an area of dried microbial mat. This dried mat extended to the waterline, where it was replaced by hydrated microbial mat. These vegetation zones are also related to the availability of water. The outer cyanobacterial zone occurred in areas where either water level reached only on rare occasions, or where winds would deposit spray from the lake and therefore provide moisture. Groundwater flow during summer could also moisten area. The inner ring of dried microbial mat 70

Figure 43: Typical zonation of moss, lichen and cyanobacterial communities in (a) stream beds and (b) around lakes. suggests that the water level in many of the lakes was low at the time of the 1999/2000 ANARE visit, but during periods of greater melt water input, water levels would increase and rehydrate this material. Antarctic microbial mats are well known to be adapted to periods of desiccation and freezing.

6. Terrestrial Fauna

In this section what little is known about the terrestrial fauna of the Bunger Hills is summarised. Most of the animals are associated with patches of moss, lichen or terrestrial algae. Also included are details of nematodes and tardigrades in algal samples from the margins of lakes.

6.1 Heterotrophic protists

Heterotrophic protists include unicellular forms, e.g., amoebae and foraminifera, which metabolise plant- or animal-derived organic material. Sudzuki (1979) gave a short list of Testacea, Amoebina, Zoomastigophora, Holotricha and Spirotricha (all heterotrophic protists) 71 found in samples collected during the 1977 ANARE visit. A list of genera recorded is given in Appendix 8. Nothing else is known at present about the occurrence of unicellular animals in the soils or lakes of the Bunger Hills.

6.2 Nematodes

Kir’yanova (1964) reported on the occurrence of nematodes in terrestrial and lacustrine samples collected during early Russian visits to the Bunger Hills. Three species, Plectus antarcticus , Plectus frigophilus and Plectus globilabiatus , were found in this material (see Appendix 9 for details). The latter two species were described from Bunger Hills samples, though the last is considered nomen dubum (Maggenti, 1961). Yeates (1979) studied 10 samples of moss and algae collected during the brief ANARE visit in 1977, and identified Plectus antarcticus in moss samples, Plectus frigophilus from moss and algae samples, and a third species, Helicotylenchus sp., from moss. Plectus globilabiatus was not observed. Nothing is known about the ecology of these nematodes. Kir’yanova (1964) also reported the occurrence of two Chromadoridae in samples collected from a marine area.

6.3 Tardigrades

Tardigrades were observed in samples collected during the early Russian visits to the Bunger Hills, but were not identified (Korotkevich, 1964c). Samples collected during the 1977 ANARE visit were found to contain at least 7 species (Barker, 1977) (Appendix 9), including the first Antarctic record of the genus Pseudechiniscus . This observation is thus of considerable biogeographical interest.

6.4 Mites and fleas

Soils samples collected from two sites during the 1977 ANARE visit contained live specimens of Nanorchestes antarcticus (Barker, 1977), which is also known from the Vestfold Hills and other ice-free areas of Antarctica. Stahle (in Ledingham 1986a) reported the widespread occurrence of mites, tentatively identified as the same species, in samples of moss and soil collected during the 1986 ANARE visit. It is possible that the Antarctic flea Glaciopsyllus antarcticus is present in Snow petrel nests (Whitehead et al., 1991), but its presence has not been reported. 72

6.5 Birds

The Bunger Hills have been described as a place of peace and calm (Markov, 1970), largely due to the lack of noisy Adélie penguin rookeries. However, birds do occur in the region; the species recorded are discussed in the following sections.

6.5.1 Snow petrel

The snow petrel ( Pagodroma nivea ) is the most common bird of the Bunger Hills, with an estimated population of 1000 breeding pairs (Bulavinstev et al., 1993) in colonies of up to 50 pairs dispersed widely throughout the southern Bunger Hills at altitudes between 25 and 155 m above sea level (Filcek and Zielinski, 1990; Verkulich and Hiller, 1994). About 70% of nests are located in cavities and cracks in bedrock on hills and ridges, while 30% breed under or between large boulders. Nest sites are chosen to be protected from snow accumulation and strong winds. A distribution map of breeding sites was published by Verkulich and Hiller (1994), which is redrawn in Figure 44. The figure shows that the sites are concentrated around the centre of the Hills, with significant numbers both along the rock-Apfel Glacier margin and the otherwise

Figure 44: Locations of nesting sites of Snow petrels (after Verkulich and Hiller, 1994: filled squares). Also shown are other nesting sites identified during the1999/2000 ANARE visit (open squares).

73 biologically depauperate areas to the north. Filcek and Zielinski (1990) observed that chicks were fed once per day at dusk or on alternate days in early February. During this period, only a small number of adults stayed in the colonies. The chicks fledged during the first week of March, and most birds had left the colonies by the middle of the month.

Snow petrels have been used indirectly in studying the Holocene deglaciation of the Bunger Hills. Verkulich and Hiller (1994) determined the age of organic deposits around nesting sites by radiocarbon techniques, and found that petrels had been breeding in the area for at least 10000 years.

Observations made during the 1999/2000 ANARE visit were consistent with those of the Russian researchers. A few extra breeding sites were identified (see Figure 44), and birds were observed to be generally more common around dusk. However, sightings of individual birds were not particularly common, suggesting direct flight between the colonies and the feeding grounds, presumably late in the evening or early in the morning.

6.5.2 Wilson’s storm petrel

Wilson’s storm petrels ( Oceanites oceanicus ) are, in the words of Filcek and Zielinski (1990), ‘…very dispersed, [with] solitary nests in the crevices of rocky rubble covering the slopes of the Hills.’ No study of the distribution or breeding of Wilson’s storm petrels has been made in the Bunger Hills, in part due to the difficulty in identifying nesting sites and the birds’ generally nocturnal and cryptic behaviour. No nesting sites were positively identified during the 1999/2000 ANARE visit, though they undoubtedly occurred. Korotkevich (1964a) included the Wilson’s storm petrel among a list of breeding birds in the Bunger Hills, and Ledingham (1986a) reported that the species was frequently sighted in the area during the 1986 ANARE visit. Birds were observed sporadically during the 1999/2000 ANARE visit at both Edgeworth David and other locations, occasionally in pairs but generally individual animals. Filcek and Zielinski (1990) recorded the last Wilson’s storm petrel for the summer on 2 March.

6.5.3 South polar skua

The south polar skua ( Catharacta mccormicki ) has been reported regularly from the Bunger Hills, where it breeds (Korotkevich, 1964a). Barker (1977) observed four skuas at his site 74

10 (Figure 4), while Ledingham (1986a) found the species to be ‘fairly common’ throughout the Hills, though absent from the Obruchev Hills to the south west. Filcek and Zielinski (1990) noted that nests of this species were very dispersed, but that the birds gathered in post-breeding groups of up to ten individual at streams or unfrozen lakes, including the outlet to Algae Lake, prior to leaving the area for winter in early April.

During the 1999/2000 ANARE visit regular sightings of skuas were made both at the field camps and when travelling on foot throughout the area. It is difficult to estimate abundance, as individual birds identifiable by, for example, missing webbing on the foot, were seen over quite large areas (of the order of ten km 2). The population was very roughly estimated to be 50 birds. The maximum number seen at one time was six (while on the sea ice south of Krylatyj Peninsula, possibly at the junction of a number of loose territories), and pairs or trios were observed regularly. Figure 45 shows the location of sightings during the 1999/2000 ANARE visit, though it should be noted that the mobility and inquisitiveness of the birds means that the sites marked on the map do not necessarily reflect true distribution or numbers. No banded birds were seen.

The absence of breeding colonies of Adélie Penguins in the Bunger Hills means that skuas must rely on snow petrels and Wilson's storm petrels for food. Remains of both these species were observed throughout the Hills, ranging in age from very fresh to mummified. The relatively small petrel population presumably limits the size of the population and breeding of the skuas, though birds may be travelling to penguin colonies near the outer edge of the Shackleton Ice Shelf to feed. Little aggressive behaviour was observed between skuas, even when ‘discussing’ a possible food item (a snow petrel carcass). On only one occasion was aggressive behaviour towards humans noted, probably indicative of the presence of a nest (for the location of this site, see Figure 45). Even in this case, the vocal and physical displays were of low intensity and soon subsided. No attempt was made to determine whether an egg or chick was actually present. Birds often perched on high points, many of which were surrounded by guano as well as regurgitated pellets containing petrel remains. In some cases lichen growth seemed to be enhanced by the nutrients provided by this material.

75

Figure 45: Location of South polar skua sightings, 1999/2000. Also shown in the location of a possible nesting site (open square). The absence of records from areas is more likely to reflect less time was spent in these areas, rather than a true absence of birds.

Skuas were attracted to field camps during habitation. Ledingham (1986a) also recorded the presence of ‘camp skuas’. The first skua to visit Edgeworth David in 1999/2000 stayed in the area for about a week, and showed little interest in plastic bags or other possible food sources around the camp. On one occasion this bird disgorged a pellet containing the remains of a Wilson's storm petrel. Towards the end of the visit, a second skua took up residence at Edgeworth David. This bird actively attacked plastic bags, presumably with the aim of getting food.

6.5.4 Adélie penguin

No breeding colony of the Adélie penguin ( Pygoscelis adeliae ) occurs within the Bunger Hills region (Filcek and Zielinski, 1990); the physical barrier of the Shackleton Ice Shelf generally prevents access for the birds to the ice-free areas. However, occasional birds do reach the area, as indicated by the observation of a live bird in the late 1950s (Korotkevich, 1964a) and the presence of mummified carcasses (Markov et al., 1970; Ledingham, 1986a). During the 1999/2000 ANARE visit a live Adélie penguin was observed and photographed on the surface of Transkriptsii Gulf offshore from Edgeworth David Station on 8 January 2000 (Figure 46). Tracks of this penguin were noted between Transkriptsii Gulf and Lake 14, indicating that it had come 76

Figure 46: The second record of a live Adélie penguin in the Bunger Hills: Transkriptsii Gulf, 8 January 2000. Edgeworth David is visible in the background. from the north. The penguin continued walking to the west from the point of observation, and was not observed subsequently.

6.6. Seals

A population of Weddell seals ( Leptonychotes weddelli ) in the marine waters of the Bunger Hills has been recorded sporadically, though no study or accurate census of this population has been reported in the literature. A picture of a Weddell Seal ‘in the Bunger Oasis’ is given in Lebedev (1959, p201), Wisniewski (1983) mentioned the population briefly, and Filcek and Zielinski (1990) reported two live animals. Live seals (circa 20), including a pup, were also observed during helicopter operations in the northern half of the Bunger Hills on the 1986 Australian visit (Ledingham, 1986a, b). Klokov et al. (1990) record the sighting of a single live seal on the sea ice in March. No live animals were observed during the 1999/2000 ANARE visit, though little time was spent along the marine coast of the study area. Evidence for this population, which is possibly isolated by the Shackleton Ice Shelf from those that occur along the coast of East Antarctica and is therefore of considerable scientific interest, was observed on the current visit in the form of five mummified carcasses in a small area of the shoreline of Rybij Khvost Gulf near Grantovaya Sopka Island. Similar mummified carcasses have been reported 77 from the area previously (Korotkevich, 1964b; Syroechkovskii and Evteev, 1964; Filcek and Zielinski, 1990). 78

Acknowledgments

The author thanks the following people who helped to make the visit to the Bunger Hills and the subsequent writing of this report a success: Belinda Harding, Garry Kuehn and Damian Flynn for assistance and comradeship in the field; James Shevlin (AAD), Mike Sharp (Polar Logistics), Doug McLeod (First Air) and Phil Howard (First Air) for arranging, supporting and piloting the Twin Otter flights to and from the Bunger Hills; Professor Warwick Vincent (Université Laval, Canada) and Jono Reeve (AAD) for lending me some equipment; Kevin Bell for supplying and defraying the costs of developing film; Rod Seppelt for discussion of the lichen and moss data; Andie Smithies and Graeme Watt for assistance with delving into the Russian and other literature in the AAD library; John Cox for assistance in preparing the base figure of the Bunger Hills. 79

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Appendix 1

Visits to the Bunger Hills: 1987-2000.

Year Beginning of Season End of Season Who Reference 1986/1987 19/1/87 15/3/87 32 nd SAE Klokov et al., 1990 1987/1988 ?1/88 ?4/88 33 rd SAE Kaup et al., 1993 1988/1989 2/89 3/89 34 rd SAE-Polish Filcek and Zielinski, 1990 1989/1990 1990/1991 ?1/91 ?4/91 36 th SAE-German Verkulich and Melles, 1992 1991/1992 20/12/91 6/4/92 37 th SAE-USA Doran et al., 1996 1992/1993 1993/1994 22/1/94 3/4/94 39 th SAE-German Melles, 1994 1994/1995 2/1/95 ANARE Antarctic Division, 12/4/95 unpublished data 1995/1996 6/12/95 23/2/96 ANARE Hudspeth, 1996 1996/1997 1997/1998 1998/1999 29/11/98 8/12/98 Polar Logistics M. Sharpe, Personal communication, 1999/2000 26/12/99 20/1/00 ANARE This report 14/3/00 ANARE G. Bain, Personal communication 88

Appendix 2

Algal and cyanobacterial species recorded in lakes of the Bunger Hills. The localities include the description from the original publication if the lake names are not known, and the sampling sites from Barker.

Species Locality Type of Benthos (B) or Reference Organisms Water Column (W) Achnanthes Lake Polest Diatom B Kaup et al., 1993 brevipes var . intermedia Amorphonstoc Transkriptsii Gulf Cyanobacterium B Vialov and paludosum Sdobnikova, 1961 Amphora sp . Small Lake, Diatom B Vialov and 5,7,10,11 Sdobnikova, 1961 Anabaena sp . 10, 11 Cyanobacterium B Barker, 1977 Chlorella sp . 5,11 Chlorophyte B Barker, 1977 Chlorella vulgaris Transkriptsii Gulf, Chlorophyte B Vialov and Small Lake Sdobnikova, 1961 Chlorococcum Algae Lake, Small Chlorophyte B Korotkevich, 1964c infusionum lake Chrococcus 7,10,11 Cyanobacterium B Barker, 1977 turgidus Chroococcus minor 10,11 Cyanobacterium B Barker, 1977 Chroococcus sp . 5, 10,11 Cyanobacterium B Barker, 1977 Coscinodiscus Lake Polest Diatom W Kaup et al., 1993 centralis Coscinodiscus Lake Polest Diatom W Kaup et al., 1993 oculus iridis Cosmarium Algae Lake Chlorophyte B Korotkevich, 1964c granatum Cosmarium sp . 5,10,11 Chlorophyte B Barker, 1977 (Desmid) Gloecapsa 10 Cyanobacterium B Barker, 1977 aeruginosa Gloecapsa Small Lake Cyanobacterium B Vialov and crepidinium Sdobnikova, 1961 89

Hantzschia sp . Algae Lake Diatom B Korotkevich, 1964c Microcystis 5,7,10 Cyanobacterium B Barker, 1977 aeruginosa Microcystis Transkriptsii Gulf Cyanobacterium B Vialov and musiciola Sdobnikova, 1961 Navicula Lake Polest Diatom B, W Kaup et al., 1993 cryptocephala var . intermedia Navicula mutica 5,7,10,11 Diatom B Barker, 1977 Navicula papula 7,10,11 Diatom B Barker, 1977 Navicula sp . Algae Lake Diatom B Korotkevich, 1964c Nitzschia sp . 5,7,10,11 Diatom B Barker, 1977 Nostoc paludosum 5,10,11 Cyanobacterium B Barker, 1977 Nostoc sp . 7,10,11 Cyanobacterium B Barker, 1977 Oocystis sp . 10, 11 Chlorophyte B Barker, 1977 Oscillatoria nigra 7,10,11 Cyanobacterium B Barker, 1977 Oscillatoria sp . 7 Cyanobacterium B Barker, 1977 Phormidium Small Lake Cyanobacterium B Vialov and bohneri Sdobnikova, 1961 Phormidium Small Lake Cyanobacterium B Vialov and favosum Sdobnikova, 1961 Phormidium retzii Transkriptsii Gulf Cyanobacterium B Vialov and Sdobnikova, 1961 Phormidium Algae Lake Cyanobacterium B Korotkevich, 1964c tenuissium Pinnularia sp Algae Lake Diatom B Korotkevich, 1964c Plectonema Transkriptsii Gulf Cyanobacterium B Vialov and battersii Sdobnikova, 1961 Plectonema Small Lake Cyanobacterium B Vialov and boryanum Sdobnikova, 1961 Schizothrix 7 Cyanobacterium B Barker, 1977 muelleri Schizothrix sp . Algae Lake Cyanobacterium B Korotkevich, 1964c Staurastrum sp . 7,10 Chlorophye B Barker, 1977 (Desmid) Stauroneis anceps 5,7,10,11 Diatom B Barker, 1977

90

Appendix 3

Organisms for which the type locality is in the Bunger Hills.

Species Type of organism Type locality Reference Acanthocyclops mirnyi Copepod Algae Lake Borutzky and Vinogradov, 1957 Grimmia doniana f. Moss South of Algae Lake Kuc, 1969 antarctica 1 Bryum korotkevicziae 2 Moss Algae Lake Savich-Lyubitskaya and Smirnova, 1964 Cnemidocarpa Ascidian Rybij Khvost Gulf? Vinogradova, 1958 zenkevitchii Dactylospora Fungus North coast of Algae Lake Olech and Alstrup, 1996 dobrowolskii Gladioferens antarcticus Copepod White Smoke Lake Bayly, 1994 Notholca verae Rotifer Algae Lake Kutikova, 1964 Paralabidocera Copepod Rybij Khvost Gulf? Brodski and Zvereva, separabalis 1976 Plectus frigophilus Nematode Freshwater pond Kir’yanova, 1964 Plectus globilabiatus 3 Nematode Moss beds on the shore of Kir’yanova, 1964 a freshwater lake

Notes:

1. Grimmia donniana is now designated Grimmia reflexidens (Ochyra, 1998). 2. After studies of material from lakes in the Vestfold Hills and the McMurdo Dry Valleys, Bryum korotkevicziae is now regarded as an aquatic form of Bryum pseudotriquetrum (Kaspar et al., 1982; Seppelt, 1983. 3. This species is considered nomen dubum by Maggenti (1961), and has not been identified in subsequent sampling. 91

Appendix 4

Rotifer species recorded in the Bunger Hills. Samples were collected from Algae Lake and other, unidentified, lakes. The samples studied by Sudzuki (1979) was from Barker’s locations 2 and 8.

Species Reference Adineta grandis Dartnall, personal communication Adineta sp. Dartnall, personal communication. Bdelloidia ? Korotkevich, 1964c; Sudzuki, 1979 Epiphanes senta Korotkevich, 1964c Habrotrocha sp. Dartnall, personal communication. Lepadella patella Korotkevich, 1964c, Dartnall, personal communication Notholca verae Kutikova, 1964a,b; Korotkevich, 1964c, Dartnall, personal communication Philodina alata Korotkevich, 1964c Philodina gregaria Dartnall, personal communication Philodina sp. Korotkevich, 1964c, Dartnall, personal communication Proales reinhardti Kutikova, 1964b Rhinoglena fertoensis Korotkevich, 1964c 92

Appendix 5

A list of fungal species recorded in the Bunger Hills.

Species Reference Acremonium strictum Barker, 1977 Actinomycete sp . Barker, 1977 Arthonia rufida Olech and Alstrup, 1996 Cladosporium cladosporoides Barker, 1977 Dactylospora dobrowolskii Olech and Alstrup, 1996 Dreschlera sp . Barker, 1977 Endococcus propinquus Olech and Alstrup, 1996 Lichenoconium usneae Olech and Alstrup, 1996 Mucor plumbens Barker, 1977 Penicillium brevi-compactum Barker, 1977 Penicillium expansum Barker, 1977 Penicillium spinulosum Barker, 1977 Penicillium spp. Barker, 1977 Penicillium verrucosum Barker, 1977 Phaeosporus usneae Olech and Alstrup, 1996 Rhizopus nigricans Barker, 1977 Thelebolus microsporus Barker, 1977

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Appendix 6.

Lichen species recorded by Barker (1977), Olech (1989) and Olech and Alstrup (1996) in studies of the botany of the Bunger Hills. Also indicated is whether the species has been recorded in the Vestfold Hills and the Windmill Islands (Casey) areas (Seppelt, 1986b, unpublished key).

Species Bunger Hills Bunger Hills Vestfold Hills Windmill (Barker, 1977) (Olech, 1989, Islands Olech and Alstrup, 1996) Acarospora gwynnii X X X X Acarospora williamsii X X Arthonia subantarctica X Buellia cladocarpiza X Buellia frigida X X X X Buellia grimmiae X X X X Buellia c.f . illaetabilis X Buellia lignoides X X Buellia pycnogonoides X X Buellia pulverulenta X Caloplaca athallina X X X Caloplaca citrina X X X Caloplaca saxicola X Candelariella flava X (as C. X (as C. X (as C. X antarctica ) hallettensis ) antarctica ) Carbonea vorticosa X Lecanora expectans X X X Lecanora polytropa X Lecidea cancriformis X X Lecidea phillipsiana X X Leprocaulon subalbicans X Physcia caesia X X X X Physcia dubia X Pleopsidium chlorophanum X X (as Biatorella cerebriformis ) Pseudephebe minuscula X (as Alectoria X X (as Alectoria X minuscula ) minuscula ) 94

Rhizocarpon geographicum X (as X X (as X Rhizocarpon Rhizocarpon flavum) flavum) Rhizoplaca melanophthalma X (as Lecanora X X (as Lecanora X melano- melano- phthalma )) phthalma ) Rinodina olivaceobrunnea X X X Rinodina petermannii X X Rinodina turfacea X Umbilicaria aprina X X X X Umbilicaria decussata X X X X Usnea antarctica X X X X Usnea sphacelata X X Xanthoria mawsonii X X (as X. X (as X. candelaria ) candelaria )) Xanthoria elegans X (as Calo- X X X placa elegans )

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Appendix 7.

Moss species recorded by Savich-Lyubitskaya and Smirnova (1964, 1966, 1971a, 1971b, 1972), Kuc (1969); Barker (1977) and Olech (1989) in studies of the botany of the Bunger Hills. Also indicated is whether the species has been recorded in the Vestfold Hills or the Windmill Islands (Casey) areas (Seppelt, 1986a; unpublished key).

Species Bunger Bunger Bunger Bunger Vestfold Windmill Hills Hills (Kuc) Hills Hills Hills Islands (Savich- (Barker) (Olech) Lyubits- kaya and Smirnova) Bryum algens X X(?) X X X (as B. pseudotri- quetrum ) Bryum X (as B. X (as B. subrotundifolium argenteum) argenteum Bryum X (as B. X (as B. X (as B. pseudotriquetrum korotkevicz inconne- korotkevicz iae) xum f. iae) toment- osum Ceratodon X X X purpureus Grimmia X (as reflexidens Grimmia doniana f. antarctica) Grimmia X plagiopodia Pohlia nutans X Hennediella heimii X (as Pott- X (as Pott- ia heimii) ia heimii) Sarconeurum X X glaciale Schistidium X(?) X X (as X (as 96 antarcticum Grimmia Grimmia antarctici ) antarctici )

Appendix 8.

Testaceae, Amoebina, Zoomastigophora, Holotricha and Spirotricha recorded by Sudzuki (1979) from terrestrial and lacustrine samples collected in the Bunger Hills. The numbers of the sample locations refer to the sites of Barker (1977) (Figure 4).

Species Class Sampling Location Amoeba sp. Amoebina 3 Anisonema sp. Zoomastigophora 2 Assulina muscorum Testacea 2,7,8 Assulina sp. Testacea 2,4 Bodo globosa Zoomastigophora 2 Bodo sp. Zoomastigophora 3,7,8,10 Bryophrya sp.? Holotricha 3 Centropyxis aerophila Testacea 10 Centropyxis sp.? Testacea 8 Chilodonella sp. Holotricha 3 Cochliopodium sp. Amoebina 7 Corythion aerophila Testacea 4,7 Corythion sp. Testacea 2,7 Difflugiella sp.? Testacea 8 Hyalodiscus sp. Testacea 7 Keronopsis sp. Spirotricha 3 Leptochlamys Testacea 7 Microcorycia briophila. Testacea 7 Microcorycia sp.? Testacea 1,8 Oxytricha sp. Spirotricha 3 Parmulina sp.? Testacea 1,3,7,8,10 Paruroleptus sp. Spirotricha 3 Psyochia sp.? Testacea 8 Spathidium breve ? Holotricha 3 Strombilidium sp. Spirotricha 3 Thecamoeba Amoebina 4 97

Trinema lineare Testacea 1 Trochilia sp.? Holotricha 3

98

Appendix 9

Nematode, tardigrade and mite species recorded in the Bunger Hills.

Species Type of organism Reference Helicotylenchus sp . Nematode Yeates, 1979 Hypsibius mertoni simoizumi Tardigrade Barker, 1977 Hypsibius sp . 1 1 Tardigrade Barker, 1977 Hypsibius sp. 2 1 Tardigrade Barker, 1977 Hypsibius? 2 Tardigrade Barker, 1977 Macrobiotus hufelandi Tardigrade Barker, 1977 Macrobiotus sp. Tardigrade Barker, 1977 Nanorchestes antarcticus Mite Barker, 1977 Plectus antarcticus Nematode Kir’yanova, 1964 Plectus frigophilus Nematode Kir’yanova, 1964 Plectus globilabiatus 3 Nematode Kir’yanova, 1964 Pseudechiniscus sp . Tardigrade Barker, 1977

Notes:

1. Barker (1977) gives no further details for these species. 2. Barker (1977) gives a photo of this species. 3. Plectus globilabiatus was considered nomen dubum by Maggenti (1961).