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Antarctic Science 1 (3): 2 15-224 (1 989) Survey of algae and other terrestrial biota at Edward VII Peninsula, Marie Byrd Land PAUL A. BROADY Department of Plant and Microbial Sciences, University of Canterbury, Private Bag. Christchurch I, New Zealand Abstract: The survey concentrated on algal communities but also gathered information on mosses, lichens andmicrofauna. All 23 nunataks in theRockefellerandAlex~dramountains(77000'-78030'S, 152"-154OW) were visited, including ones both with and without nutrient enrichment from bird colonies. Over 30 species of algae were recorded with Cyanobactena and Chlorophyta dominant as in other regions of continental Antarctica. Diatoms were not found. The most frequent algae were Cyanothece aeruginosa, Gloeocapsa spp., Oscillatoriaceae, Nostoc sp., Pseudococcomyxa simplex, Stichococcus bacillaris, cf. Desmococcus vulgaris, Prasiola crispa and Prasiococcus caicarius. Although lichens were the most visually prominent vegetation,free-livingalgae occurredin the widespread 'non-aquatic' habitatsas epilithic,chasmoendolithic andedaphic communitiesas well as being epiphytic on the sparse moss cushions. Aquatic habitats were few and there were only three smallpondson or adjacent tonunataks. On ablating ice, cryoconitepondscontained Homoeothrix cf. rivularis, recorded for the first time in Antarctica. Preliminary identifications show 23 speciesof lichens and six species of mosses. No mites and collembola were seen. Samplescontained testate amoebae, ciliate protozoa, rotifers and tardigrades but no nematodes. Received 30 December 1988, accepted 10 March 1989 Key words: Antarctica, ecology, microfauna, nunataks, vegetation. Introduction The majority of terrestrial biological research in continental Antarctica has been performed at coastal ice-free regions with good access from research stations, for instance in the dry valleys of southern Victoria Land and on Ross Island (e.g. Friedmann 1982, Broady 1989), at the Vestfold Hills (Pickard 1986)andin thevicinity of Syowa station, Liitzow- Holm Bay (e.g. Oguni et al. 1987). However, there is increasing interest in the more remote nunataks, e.g. Mount Kobourg, northern Victoria Land (Kappen 1985) and in Dronning Maud Land (Engelskjon 1986, Pankow et al. 1987, Ryan & Watkins, in press). Wide-ranging observations and collections were made during a combined geological and biological expedition to nunataks on Edward VII Peninsula (77°00-78"30S, 152"- 154"W),Marie ByrdLand (Fig. 1) during summer 1987-88. Reported here are observations on the algae, mosses,lichens and microfauna. A description of the ornithology has been published elsewhere (Broady et a/. 1989). Reportson the terrestrialbiology of Edward VII Peninsula are few and lack detail. Early reports of mosses and lichens were made by Presuud (in Amundsen 1912,p. 249), Gould (1931, p. 20) and Siple (1938, p. 490). The last author reported mainly on more detailed studies made in Edsel Ford Ranges, 130 km further east (77"S, 145"W). Lichens were described by Dodge & Baker (1938) and mosses by Baruam (1938). Perkins(1945,p. 282) sampledplants from 11peaks in RockefellerMountains. Lichens in these collectionswere 1 Fig. 1. Location of Edward VII Peninsula and other regions mentioned in the text. 1-3 = Marie Byrd Land, 1 = Edward VII Peninsula, 2 = Edsel Ford Ranges, 3 = Ames and Flood ranges, 4 = Horlick Mountains, 5 = Ellsworth Mountains, 6-8 = Dronning Maud Land, 6 = Robertskollen, 7 = Gjelsvikfjella and Miihlig-Hofmannfjella, 8 = Unterseeoase, 9 =Liitzow-Holm Bay, 10 = Mawson Rock, 11 = Framnes Mountains, 12 = Vestfold Hills, 13-14 = northern Victoria Land, 13 = Mount Kobourg, Birthday Ridge, 14 = Edmondson Point, Cape Washington, Campbell Glacier, Inexpressible Island, 15 = southern Victoria Land dry valleys, 16 = McMurdo Ice Shelf, 17 = Ross Island, 18 = Shackleton Glacier. 21 5 https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0954102089000337 Downloaded from https://www.cambridge.org/core. IP address: 54.39.17.49, on 01 Apr 2018 at 13:19:39, subject to the Cambridge Core terms of use, available at
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Page 1: Survey of algae and other terrestrial biota at Edward VII Peninsula ...

Antarctic Science 1 (3): 2 15-224 (1 989)

Survey of algae and other terrestrial biota at Edward VII Peninsula, Marie Byrd Land

PAUL A. BROADY Department of Plant and Microbial Sciences, University of Canterbury, Private Bag. Christchurch I , New Zealand

Abstract: The survey concentrated on algal communities but also gathered information on mosses, lichens andmicrofauna. All 23 nunataks in theRockefellerandAlex~dramountains(77000'-78030'S, 152"-154OW) were visited, including ones both with and without nutrient enrichment from bird colonies. Over 30 species of algae were recorded with Cyanobactena and Chlorophyta dominant as in other regions of continental Antarctica. Diatoms were not found. The most frequent algae were Cyanothece aeruginosa, Gloeocapsa spp., Oscillatoriaceae, Nostoc sp., Pseudococcomyxa simplex, Stichococcus bacillaris, cf. Desmococcus vulgaris, Prasiola crispa and Prasiococcus caicarius. Although lichens were the most visually prominent vegetation, free-living algae occurred in the widespread 'non-aquatic' habitats as epilithic, chasmoendoli thic andedaphic communities as well as being epiphytic on the sparse moss cushions. Aquatic habitats were few and there were only three small pondson or adjacent tonunataks. On ablating ice, cryoconite pondscontained Homoeothrix cf. rivularis, recorded for the first time in Antarctica. Preliminary identifications show 23 species of lichens and six species of mosses. No mites and collembola were seen. Samples contained testate amoebae, ciliate protozoa, rotifers and tardigrades but no nematodes.

Received 30 December 1988, accepted 10 March 1989

Key words: Antarctica, ecology, microfauna, nunataks, vegetation.

Introduction

The majority of terrestrial biological research in continental Antarctica has been performed at coastal ice-free regions with good access from research stations, for instance in the dry valleys of southern Victoria Land and on Ross Island (e.g. Friedmann 1982, Broady 1989), at the Vestfold Hills (Pickard 1986) andin thevicinity of Syowa station, Liitzow- Holm Bay (e.g. Oguni et al. 1987). However, there is increasing interest in the more remote nunataks, e.g. Mount Kobourg, northern Victoria Land (Kappen 1985) and in Dronning Maud Land (Engelskjon 1986, Pankow et al. 1987, Ryan & Watkins, in press).

Wide-ranging observations and collections were made during a combined geological and biological expedition to nunataks on Edward VII Peninsula (77°00-78"30S, 152"- 154"W), Marie ByrdLand (Fig. 1) during summer 1987-88. Reported here are observations on the algae, mosses, lichens and microfauna. A description of the ornithology has been published elsewhere (Broady et a/. 1989).

Reports on the terrestrial biology of Edward VII Peninsula are few and lack detail. Early reports of mosses and lichens were made by Presuud (in Amundsen 1912, p. 249), Gould (1931, p. 20) and Siple (1938, p. 490). The last author reported mainly on more detailed studies made in Edsel Ford Ranges, 130 km further east (77"S, 145"W). Lichens were described by Dodge & Baker (1938) and mosses by Baruam (1938). Perkins(1945,p. 282) sampledplants from 11 peaks in Rockefeller Mountains. Lichens in these collections were

1 Fig. 1. Location of Edward VII Peninsula and other regions

mentioned in the text. 1-3 = Marie Byrd Land, 1 = Edward VII Peninsula, 2 = Edsel Ford Ranges, 3 = Ames and Flood ranges, 4 = Horlick Mountains, 5 = Ellsworth Mountains, 6-8 = Dronning Maud Land, 6 = Robertskollen, 7 = Gjelsvikfjella and Miihlig-Hofmannfjella, 8 = Unterseeoase, 9 =Liitzow-Holm Bay, 10 = Mawson Rock, 11 = Framnes Mountains, 12 = Vestfold Hills, 13-14 = northern Victoria Land, 13 = Mount Kobourg, Birthday Ridge, 14 = Edmondson Point, Cape Washington, Campbell Glacier, Inexpressible Island, 15 = southern Victoria Land dry valleys, 16 = McMurdo Ice Shelf, 17 = Ross Island, 18 = Shackleton Glacier.

21 5

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21 6 P.A. BROADY

identified by Dodge (1973). Rudolph (1967) collected lichens and noted an absence of microarthropods on Washington Ridge. Thisis theonly commenton microfauna although in eastern Marie Byrd Land, Siple (1938, p. 495) found protozoa, rotifers and tardigrades in pond water, and Strandtmann (1978,1981) described a new species of mite.

Description of the region

The seven nunataks of the Alexandra Mountains and the 16 of theRockefeller Mountains are the only ice-free land in the Edward VII Peninsula, and comprise less than 0.5% of the total area (Fig. 2). They are typical nunataks of the 'Conti- nental Ice Slope' region as defined by Pickard & Seppelt (1984, p. 88, fig. 10). There are no coastal outcrops. Scott Nunataks are closest to the coast and are 11 km inland. Mount Paterson is furthest inland, 86 km from the coast. Summits range from about 419 m to 1174 m altitude and, except close to the coast, the surrounding ice-surfaces have an elevation of 400 m to over 1000 m. The most extensive nunataks, Mount Paterson and Washington Ridge in Rockefeller Mountains (Figs 3-5), are low ridges up to 3 km long. Ice-free rock in Alexandra Mountains is much less extensive and is mostly steep slopes and cliffs of two N-NE facing escarpments. The rocks of both ranges comprise

R O S S

S E A

77'30' E O W A R O V I I 0 +q-& .\MA

C'L 3 i P E N I N S U L A

i R O C K E F E L L E R Ills

k m

906.

Fig. 2. Geographical features of Edward VII Peninsula, Mane Byrd Land. Inset shows the location of Edward VII Peninsula in the south-east comer of Ross Sea. Nunataks of the Alexandra and Rockefeller mountains are shown as follows: SN = Scott Nunataks, SW = Mount Swadener, BO = Bowman Peak, MA = Mount Manger, JO = Mount Josephine, CL = Clarke Peak, LG = La Gorce Peak, DR = Drummond Peak, FR = Mount Frazier, JA = Mount Jackling, FZ = Mount Fitzsimmons, SH = Mount Shideler, PA = Mount Paterson (see Fig. 4), GO = Gould Peak, BU = Mount Butler, TE = Tennant Peak. Nunataks to the east of Mount Paterson are shown in more detail in Fig. 3.

granites and metasediments. A general description of the region has been provided by Wade (1945) and a 1 :250 000 map is available (United States Geological Survey 1972).

On 47 days in the field, during early to mid-summer, meteorological observations were taken twice a day at approximately 1000 and 2230 h solar time. The air tempera- ture ranged from -1 7" to -2°C but on 82% of occasions was between -10" to -3°C. There were 6-8 oktas of cloud on 76% of observations. Wind was frequent but did not exceed 20 kn. Conditions were calm for 13% of observations, winds were <10 kn for 68% and >10 kn for 19%. The prevailing wind was from east to north-east. Snowfall or snowdrift occurred on 25 days.

Methods and materials

Exposed rock was reached at all nunataks except Mount Manger where it was inaccessible. Efforts were made to collect the full diversity of vegetation at each nunatak. Samples of algae and substratum were collected into sterile polythene bags using clean instruments. In the field, speci- mens were kept at ambient temperatures and were buried in snow at campsites. Field searches were made for mites and collembola using a lox hand-lens in likely habitats but no extraction procedures were used. Transport to New Zealand and subsequent storage was at -20°C.

Taxonomy of cyanobacteria follows the system of Geitier (1932). Identifications of ciliate protozoa have been made by Dr W. Foissner (Universiat Salzburg). Rotifers and

M t L B l R T ROCKS t11.45 S l R I O t R ROCKS

MT PATERSON

4.5.6.19.20.37

\ Ill SCHLOSSBhCH 22 50 PETREL

2.39 k23,38 4 46, FOKKLR ROCKS

AE[KINRIDGt PK \ b

Wr FRANKLIN ' q5' WASHINGTON

RIDGE

48,L9, 4 M E L T /d*30

N FLOW I

0 k m 2 h L__

LAKE

Fig. 3. Nunataks in the central Rockefeller Mountains, Mount Paterson and Washington Ridge are shown in Figs 4 and 5 respectively. Proglacial lakes and position of the Antarctic petrel colony are indicated. Inset gives location (also see Fig. 2) . Numbers refer to samples (see Tables i1-IV).

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MARIE BYRD LAND ALGAE AND OTHER TERRESTRIAL BIOTA 21 7

Fig. 4. An aerial view of Mount Paterson taken from the north- west at an altitude of about 1000 m. The nunatak is about 2.8 k m lone.

Fig. 5. A view looking north along Washington Ridge from the southernmost rock exposure. A frozen pond, which is approximately 10 m long, can be seen in a windscoop in the middle distance.

tardigrades observed during microscopic examination of samples were identified by A. Suren (University of Canter- bury). Preliminary identifications of lichens and bryophytes were made by Dr R. Seppelt (Australian Antarctic Division) and Dr R.I. Lewis Smith (British Antarctic Survey).

Table I. Algae obselved in microscopic examination of 53 samples and subjective estimates of their abundance.

9% of occurrence and

rare occ. freq. abund. TG Algae abundance in all samples

-

Cyanobacteria Aphanocapsa eiachista West & West Chroococcus minor (Kuetz.) Naeg. Chroococcus turgidus (Kuetz.) Naeg. Cyanothece aeruginosa (Naeg.) Komarek Gloeocapsa aipina Naeg. Gioeocapsa kuetzingiana Naeg. Gioeocapsa cf. punctata Naeg. Gloeocapsa ralfsiana Kuetz. cf. Chroococcidiopsis sp. Crinalium cf. magnum Fritsch & John Oscillatoriaceae, trichomes 1-2 pm wide Phormidium autumnale (Ag.) Gom. Phormldium cf.fragiie (Menegh.) Gom. Phormidium cf. laminosum Gom. Schizothrir cf. antarctica Fritsch Calothrir cf. parietina (Naeg.) Thuret Homoeothrix cf. rivularis (Hansg.)

Nostoc sp. Toiypothrix cf. bouteiilei (Breb. &

Stigonema minutum (Ag.) Hassall

Komarek & Kann

Desm.) Forti

Chlorophyta cf. Chioreiia SQ.

cf. Chlorococcum sp. cf. Coenocystis sp. cf. Dicfyochloropsis sp. Pseudococcomyxa simplex (Mainx) Fott Unidentified unicells Lichen gonidia with unicells Stichococcus baciilaris Naeg. cf. Desmococcus vulgaris Brand Prasioia caiophyila (Carmich.) Menegh. Prasiola crispa (Lightf.) Menegh. Prasioia uniseriate filaments Prasiococcus caicarius (Boye Pet.)

Vischer

Chrysophyta cf. Botrydiopsis sp.

1 1 3 2

1 2

1 2 2

1

1

1 1

1 1

2

5 2

1

1 1 2 1 1 1

2 1 1

3

1

2

4 3 1 3 3

3

1

4

3

2

1

4

3

1

2

1 1

3 5

11 5

1

4

2 2 2

2 19 1 8 1 11 2 8 1 11 1 4

2 1 6

9 2

3 15 1 2

8

3 6 5 19

4 2 6

4 2

3 13 2 6 5 17 1 17 4 19 3 42

14 45 2 4 5 1 1 1 8

3 13

2

Results and discussion

Algae

The algal flora of over 30 species (Table I) could probably be extended, especially for chlorophytes, by using cultures. Several of the present identifications are tentative and require cultures for confirmation. The algal flora of comparable ‘terrestrial habitats’ at Vestfold Hills has at least 77 species, of which 45 were found by microscopic examination of samples and the remainder from cultures (Broady 1986, table 6.5).

The flora, dominated by cyanobacteria and chlorophytes, is similar to that of the Vestfold Hills where 47% and 36% of all species were cyanobacteria and chlorophytes respect- ively. Surprisingly, only a single chrysophyte was observed and no diatoms were found as either living cells or empty frustules. This contrasts with other regions where diatoms are usually found in both aquatic and non-aquatic habitats, e.g. ponds on Ross Island (Broady 1989, table 1) and ‘non- aquatic’ habitats at Vestfold Hills (Broady 1986, table 6.5) and nunataks (Pankow et al. 1987, p. 72, Ryan & Watkins, in press). Although Engelskj@n( 1986) does not mention dia- toms on nunataks in Dronning Maud Land they were present

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21 8 P.A. BROADY

(O.M. Skulberg & T. Engelskjon, personal communication 1988).

Three cyanobacteria are of particular interest. Homoeothrix has been recorded only once before from the Antarctic continent (Broady 1981a, p. 266, fig. 7). Specimens from Edward VII Peninsula closely resemble H . rivularis KomArck & Kann 1973 (p. 206, figs 3847). There are only two previous records of Stigonema minutum (Broady 1986, p. 189, Ohtani & Kanda 1987, p. 261). Cyanothece aeruginosa has been recorded frequently from Antarctica (e.g. Broady 1986, Pankow et al. 1987, Ohtani & Kanda 1987), however, the occurrence of two size forms as suspected by Komkek (1 976, p. 150) in material from Europe has not been confirmed before now. Measurements made on cells from sample 29 (Table 111) clearly demonstrated the occurrence of two populations centred around cell widths of about 10 pm and and 16 pm.

In the following sections algal communities are described from the full range of habitats at Edward VII Peninsula and are compared with those elsewhere in Antarctica. Algae in individual samples are listed in Tables 11-IV and locations of samples are shown in Figs 2 and 3.

Epilirhic algae

Thin black crusts of algae were observed infrequently where melt-water trickled over the rock surfaces. In the Alexandra Mountains black streaks down inaccessible cliff-faces were seen at Mount Bowman, Mount Swadener and Scott Nunataks. Samples from Mount Franklin and Breckinridge Peak in Rockefeller Mountains (Table 11, samples 1 and 2) both contained abundant Gloeocapsa spp. The former was unusual in having as co-dominants Homoeothrix cf. rivularis and Stigonema minutum.

Black epilithic crusts have a variable occurrence in coastal continental Antarctica. They were not found at Edmondson Point, Cape Washington and Inexpressible Island in northern Victoria Land (Broady 1987a) but did occur at Vestfold Hills and Mawson Rock (Broady 1981~). At these locations and in Dronning Maud Land (Engelskjon 1986, Ryan & Watkins, in press) Gloeocapsa spp. were dominant as at Rockefeller Mountains. However, Homoeothrix sp. has been recorded previously only at Mawson Rock (Broady 1 98 1 a).

Although absent at Alexandra Mountains, at Rockefeller

Table 11. Species composition and estimates of abundance for epilithic and chasmoendolithic algal communities and associated microfauna.

Algae FA'

1

Cyanobacteria Cyanothece aeruginosa Gloeocapsa alpina Gloeocapsa kuetzingiana a Cioeocapsa cf. punctata Gioeocapsa ralfsiana a cf. Chroococcidiopsis Oscillatoriaceae, trichomes

Phormidiwn auiumnale Phormldium cf. laminosum Calothrix cf. parietina Homoeothrix cf. rivularis a Nostoc sp. Sfigonema minutum a

Chlorophyta cf. Coenocystis sp. Pseudococcomym simplex Unidentified unicells Lichen gonidia with unicells Stichococcus bacillaris cf. Desmococcus vulgaris

Microfauna Rotifers

1-2 pm wide

BR 2 -

a f

r

0

f

Sample material, locatlon and number Epilithic Chasmoendolithic

Rockefeller Mountains Rockefeller Mountains Alexandra Mountains DR NI NI NI NI FA DR DR ST PA JA FR LG SW SN SN 3 4 5 6 1 8 9 10 11 12 13 14 15 16 17 18

0

a 0

a f f a f f f f

+ + +

0

0

f 0

a f

a

a

f a

f f a

f f + r o a

a f

a a f f a a a

I Nunataks: FA = Mount Franklin, BR = Breckinridge Peak, DR = Dmmmond Peak, NI = Mount Nilsen, ST = Strider Rocks, PA = Mount Paterson, JA = Mount Jackling, FR = Mount Frazier, LG =La Gorce Peak, SW = Mount Swadener, SN = Scott Nunataks. Subjective estimate of abundance: I = rare, o = occasional, f = frequent, a = abundant..

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MARIE BYRD LAND ALGAE AND OTHER TERRESTRIAL BIOTA 21 9

Mountains green algal crusts and associated lichens on stones on scree slopes were much more frequent than black crusts. These were usually around the bases of nunataks where exposed rock met surrounding icefields, Siple (1938, p. 490) had noted green algae in this zone at Washington Ridge. They also occurred where melt-water percolated downslope, mostly on the shaded and relatively moist lowcr sides and undersurfaces of the superficial layer of stones.

All six samples (Table 11, samples 3-8) were either domi- nated by lichens or had lichens as a major component. The lichens had enhanced development of the unicellular green phycobiont and superficially appeared to be green crusts of free-living algae. Associated free-living algae were predominantly Stichococcus bacillaris, cf. Desmococcus vulgaris and Pseudococcomyxa simplex.

Similar green algal crusts have been described at Cape Washington, northern Victoria Land (Broady 1987a, p. 12) and on Ross Island (Broady 1989). However, at both those coastal locations the crusts were dominated by free-living chaetophoralean algae, including specimens resembling cf. D. vulgaris of this study.

Chasmoendolithic algae

A striking feature at granitic nunataks was the abundance of chasmoendolithic lichens and less frequently of free-living algae (Table 11, samples 9-18). The former grew in narrow cracks between and below translucent crystals of quartzite and feldspar. They were mostly absent from opaque metasediments except where these contained veins of quartzite as on Drummond Peak and La Gorce Peak. The free-living algae usually formed green crusts and cf. D. vulgaris was often dominant. Cyanobacteria were infrequent but Nostoc sp., Gloeocapsa cf. punctata and cf. Chroococcidiopsis sp. were each abundant in a single sample.

Endolithic algae were scarce at Robertskollen due to unsuitable rock types (Ryan & Watkins, in press) however, ‘Pleurococcus’ was frequent at more easterly nunataks (Engelskjfln 1986, p. 217). This group includes species similar to cf. D. vulgaris and to Desmococcus sp. A, a chasmoendolith at Mawson Rock and nearby nunataks in North Masson Range and at Vestfold Hills (Broady 19816, p. 263, table 1). At coastal Mawson Rock, Desmococcus sp. A occurred only in areas not swept by sea-spray. In areas receiving salineaerosols P. calcarius was dominant (Broady 1981b, table 2). The generally low salt environment at inland nunataks appears ideal for the growth of cf. D. vulgaris and similar species. However, the dominant endo- lithic algae at these locations are in contrast to those at upland sites with similarly low salt levels in southern Victoria Land where a unicellular chlorophyte, Hemichloris antarctica Tschermak-Woess & Friedmann, is often accompanied by coccoid cyanobacteria (Friedmann & Ocampo-Friedmann 1984, p. 180). This difference could be related to differing

rock types or to lower moisture availability in that high altitude desert.

Soil algae

Soils were either raw mineral soils (lithosols), or nutrient- enriched (ornithogenic) in which there was a substantial organic component derived from bird guano, feathers and carcasses. Lithosols were very restricted in extent and occurred as small areas of mineral fines on sorted ground, as deposits in crevices and on ledges, and as coarse gravelly products of decomposition of the more friable granitic rocks. Ornithogenic soils occurred in the vicinity of Antarctic petrel and snow petrel nests on Washington Ridge and Mount Paterson in Rockefeller Mountains and amongst abandoned nesting sites (Broady et at. 1989) on the lowest rocky knoll of Scott Nunataks.

Lichens and algae were occasionally visible as thin black crusts over lithosols. The most frequent algal components (Table 111, samples 21-25) were C. aeruginosa, cf. D. vulgaris and S. bacillaris; several other algae were abundant in single samples. Green mucilaginous colonies of cf. Coenocyslis sp. amongst gravel and small stones were encountered twice (Table 111, samples 19 and 28). Water- flushed soils were rare but on Mount Fitzsimmons and Mount Jackling narrow gullies containing wet gravel supported ribbon-like thalli of Prasiola calophylla (Table 111, samples 26 and 27).

Foliaceous Prasiola crispa occurred downslope from the large colony of Antarctic petrels on the southernmost out- crop of Mount Paterson (Fig. 3; Table 111, samples 31 and 32). Mats covered approximately 500-1000 m2 where the ground received melt from adjacent snow drifts. The only other occurrence of P. crispa was at Scott Nunataks (Table 111, samples 33, 35 and 36) near the abandoned nest sites. Frequent microscopic associates were cf. D. vulgaris and P. calcarius. Also, these two species dominated the green crusts on a gravel soil close to the nesting sites of snow petrels on Washington Ridge (Table 111, sample 30).

Algae populations on lithosols were small in comparison with coastal regions of Antarctica, e.g. Vestfold Hills (Broady 1986,~. 18l)andRossIsland (Broady 1989) wheresaturated ground is often covered with mats of Oscillatoriaceae and mucilaginous colonies of Nostoc. The cause could be the absence of poorly drained substrata as most slopes were covered with boulders and well-drained stoney ground. At Gjelsvikfjella and Miihlig-Hofmannfjella there was more extensive wet ground with mats of these cyanobacteria (Engelskjfln 1986, table 2, p. 217) .

No other study h a s recorded such a high frequency of C. aeruginosa in lithosols despite its occurrence in several regions, both coastal (Broady 1986, table 6.5,1987a, table 1, Ohtani & Kanda 1987, p. 259) and at inland nunataks (Pankow et al. 1987, p. 69, Ryan & Watkins, in press). In

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220 P.A. BROADY

Table 111. Species composition and estimates of abundance for soil algae communities and associated microfauna.

Sample material, location and number

'NI NI FR SC FO GO FZ FZ JA LG SN WA PA PA SN SN Sh' SN 19 20 21 22 23 24 25 26 21 28 29 30 31 32 33 34 35 36

Bird-influenced Algae Lithosols

Cyanobacteria Chroococcus minor r Cyanothece aeruginosa o f f r a a Gloeocapsa kuetzingiana f 0 f Gloeocapsa cf. punctata r Gloeocapsa ralfsiana f Oscillatoriaceae, trichomes

Phormidium auumnale 0 r f Phormrdium cf. laminosum f a Schizothrir cf. antarctica a Calothrix cf. parietina 0

Nostoc sp. r a Stigonema minutum a r

1-2 um wide a

Chlorophyta cf. Chiorella sp. cf. Coenocystis sp. Pseudococcomyxa simplex Unidentified unicells Stichococcus bacillaris cf. Desmococcus vulgark Prasiola calophylla Prasiola crispa Prasiococcus calcarius

a f

a 0

0 0 a 0

r o r f o f r r a o r a a a a a f a a

a a a a a . f a a f f

a a

Chrysophyta Botrydiopsis s p

Microfauna Testate amoebae Vorticella sp. Ciliate protozoa Rotifers + Tardigrades +

0

+ +

+ + + + + + + + + + +

+

+

' Symbols as for Table I1 and SC = Mount Schlossbach, FO = Fokker Rocks, GO = Mount Gould, FZ = Mount Fitzsimmons, WA = Washington Ridge.

contrast is the lack of diatoms, as elsewhere these are common, especially Hantzschia amphioxys (Ehr.) Grun. and Navicula muticopsisvan Heurck, e.g. Vestfold Hills (Broady 1986, table 6.4), and southern Victoria Land (Seaburg et al. 1979, table 2). N. muticopsis can also be abundant amongst P. crispa on bird-influenced soils, e.g. Vestfold Hills (Broady 1986, p. 183) and Inexpressible Island (Broady 1987a, p. 13), and its absence at the Mount Paterson bird colony is unusual. P. crispa iscommon near birdcolonies on nunataks (Engelskjen 1986, p. 218, Ryan & Watkins, in press).

The occurrence of P . calcarius on bird-influenced soils is probably related to high salt levels rather than to nutrient- enrichment. The preference of this alga for saline environments without bird-influence has been noted in Europe (Lund 1966, p. 307) and Antarctica (Broady 1983, p. 216). More recently it has been found on ground close to penguin rookeries in northern Victoria Land (Broady 1987a, p. 13)

and on Ross Island (Broady 1989).

Algae epiphytic on moss

Moss cushions were frequently covered with black crusts of lichens and algae. The epiphytic algal flora (Table IV, samples 37-44) was dominated by Nostoc sp., C. aeruginosa and P. simplex. Moss cushions invariably bear crusts of epiphytic algae and the flora recorded here is not unusual (cf. Ohtani & Kanda 1987) except for the absencc of dia- toms. Although rarely dominant, diatoms are usually fre- quent, especially Pinnufaria borealis Ehr. (Broady 1982, p. 222, 1986, p. table 6.4, Ohtani 1986, p. 218). P . simplex appears to favour moss as a substratum at Edward VII Peninsulaas it does elsewhere in Antarctica (Broady 1987b, table 1).

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MARIE BYRD LAND ALGAE AND OTHER TERRESTRIAL BIOTA 221

Table Iv. Species composition and estimates of abundance for algal communities amongst bryophytes and in ponds and other samples, and associated microfauna.

~~ ~ ~ ~

Sample material, location and number

NI' FO BR BU SN SN SN S N ST FO PA WA WA PA FA NI SU 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53

Algae Bryophytes Ponds Other

Cyanobacteria Aphanocapsa elachista Chroococcus turgidus Cyanothece aeruginosa Gloeocapsa alpina Gloeocapsa kuetzingiana Gloeocapsa cf. punctata Gloeocapsa ra[fsiana Crinaliwn cf. magnum Oscillatoriaceae, trichomes

Phormidium autumnale Phormidium cf. fragile Phormldiwn cf. laminosum Calothrix cf. parietina Homoeothrix cf. rivularis

Tolypothrix cf. bouteillei

Chlorophyta cf. Chlorella sp. cf. Chlorococcum sp. cf. Coenocystis sp. cf. Dictyochloropsis sp. Pseudococcomyxa simplex a Unidentified unicells Lichen gonidia with unicells o Stichococcus bacillaris cf. Desmococcw vulgaris Prasiola crispa Prasiola crispa uniseriate

filaments Prasiococcus calcarius

1-2 pm wide

NOSfOC sp. 0

Microfauna Testate amoebae + Vorticella sp. Rotifers Tardigrades +

r f r

a

r 0

0 a

f f

r

a f

+

0

1

0

f

a f o f

a

a

a a f

f

f 0 f f r

f

a a f

a f f f a a o

' Symbols as for Table I1 and BU = Mount Butler.

a a f

+ +

+ + + + + +

Pond algae

Aquatic habitats were rarely encountered. Small but com- pletely frozen ponds in windscoops were seen on Mount Paterson, Washington Ridge and Mount Jackling but were absent from Alexandra Mountains. Other than these, only two ponds were encountered on ice-free ground, both in Rockefeller Mountains (Table IV, samples 45 and 46). One was a 10 cm deep pool in a granite boulder at Strider Rocks in which the only algae were Gloeocapsa ralfsiana and S. bacillaris. The other was a nutrient-enriched pool close to nesting skuas on the middle outcrop of Fokker Rocks. This contained rich red-brown felts of Phormidium cf. laminosum.

Another much larger nutrient-enrichedpond occurred on ice at the base of the Antarctic petrel rookery on Mount Paterson (Fig. 3, Table IV, sample 47) and contained abundant uniseriate P. crispa and felts of P. cf. faminosum. The algal floraofthesenutrient-enrichedponds is not unusual although it differs from the dense populations of Chlamydomonas sp. in a petrel rookery pool in Muhlig-Hofmannfjella (Engelskjon 1986, p. 217) and in penguin rookery pools on Ross Island (Broady 1989, table 3).

The most extensive areas of open water were two shallow lakes in ice basins 1.5 and 0.5 km south-west of Washington Ridge and Mount Paterson respectively (Fig. 3). At the time of our visit to Washington Ridge, substantial melt-water was

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222 P.A. BROADY

pouring into the lake, c. 300 m wide, from a wide area of ablating ice which had received wind-blown sand and gravel blown off the nunatak. The cryoconite ponds in this area were the most extensive freshwater habitat containing visible algae (Table IV, samples 48-50). Abundant, small, dark brown wefts of H . cf. rivularis, cf. Dictyochforopsis sp. and S . bacillaris filaments occurred amongst the mineral material. Identical, although less extensive pools were found south- west of Mount Paterson adjacent to a line of moraine. The flora was similar except for the absence of H . cf. rivularis and the presence of Nostoc sp.

Homoeothrix has not been observed previously in cryoconite pols, although Nostoc and Oscillatoriaceae are common in other regions (Wharton et al. 1981, Broady 1987a, table 1). The absence of diatoms is unusual as these are often abundant elsewhere. Pinnularia cymatopleura West & West is particularly characteristic of this habitat, e.g. on McMurdo Ice Shelf (Kellogg & Kellogg 1987, p. 85), Ross Island (Broady 1989, table 1) and on Campbell Glacier, northern Victoria Land (Broady 1987a, p. 15).

Algae porn other habitats

Occasionally bird bones were found supporting green algal crusts dominated by S. bacillaris, cf. D. vulgaris and P. calcarius (Table IV, samples 52 and 53). The latter two algae were common in this niche at Vestfold Hills (Broady 1986, p. 184, figs 28, 29). P. calcarius also formed rich crusts over the surface of an old cache of dog food left by the Second Byrd Expedition of 1933-35 at the northern end of Mount Franklin (Table IV, sample 41).

Lichens

All nunataks had lichens; overall they were the most abundant vegetation. Species listed (Table V) represent a preliminary examination of only a small proportion of the large number of specimens, which clearly contain other as yet unidentified species. The most prominent species were Pseudephebe minuscula, Umbilicaria decussata and Usnea sphacelata, with Caloplaca spp. and Xanthoria spp. near nesting birds.

Thedominant speciesresemble thoseof GroupEdescribed by Pickard & Seppelt (1984, table 3, p. 94). This community grows close to the polar ice sheet where there is relatively high moistureavailability and little wind-blown salt. Similar communities were described by Kappen (1985) in northern Victoria Land although Usnea was very rare on Mount Kobourg, the inland nunatak, but abundant at coastal Birthday Ridge.

Excellent descriptive observations on the ecology of lichens in Marie Byrd Land have been made by Siple (1938) and little can be added to these. Locally, water availability was a major factor limiting lichen growth. Epilithic lichens grew

in abundance only where they would receive moisture from melting snowdrifts, for instance on ledges, in depressions and amongst boulders on scree slopes e.g. at Drummond Peak where numerous small snowdrifts were scattered over the boulder slopes and were the only water source for the abundant epilithic lichens. Lithosols also had black crusts of lichens, especially where the soils were in slight de- pressions attracting drift snow.

Abundant chasmoendolithic lichens were present in coarse- grained granites containing large feldspar crystals, e.g. at Mount Frazier, Mount Jackling, Mount Fitzsimmons and Mount Shideler in northem Rockefeller Mountains. Epilithic lichens were often sparse on these more rapidly weathering granites whereas richer growths occurred on more stable micro-granites and in particular on schists, e.g. on Mount Nilsen, Breckinridge Peak, Drummond Peak, Clarke Peak and Scott Nunataks.

Abundant growths of large lichen thalli were present immediately adjacent to the ice edge at Mount Frazier, Mount Nilsen, Tennant Peak and Breckinridge Peak in Rockefeller Mountains. This suggests that there has been no major ice retreat in recent years. Although Siple (1938, p.

Table V. A preliminary list of lichens and bryophytes from Edward VII Peninsula, Marie Byrd Land

Lichens Biatorellu cerebriformk Filson

Buellia frigido Darb. cf. Bueliia soredians Filson Buellia sp. Caloplaca athallina Darb. Caloplaca sp. Candelariella hallettensis (Murray) Ovst.

Lecidea physciellu Darb. Lecidea sp. cf. Lecanora sp. cf. Lepraria sp. Physcia cuesia (Hoffm.) Hampe Pseudephebe minuscula (Nyl. ex Amold) Brodo & Hawksw. Pseudephebe aff. pubescens 6.) Brodo & Hawksw. Rhizocarponflavum Dodge & Baker Rhizocarponflavum f. subfoliosum Filson Rhizoplaca melanophthalma (Ram.) Lwck. & Poelt Rinodinu olivaceobrunnea Dodge & Baker Umbilicaria aprina Nyl. Umbilicariu decursata (Vill.) Zahlbr. Usnea sphacelata R. Br. Xanthoria elegans (Link.) Th. Fr. Xanthoria muwsonii Dodge

Mosses Bryum argenteum Hedw. Bryum pseudotriquetrum (Hedw.) Schwaegr. Grimmia antarcfici Card. Grimmia lawiana Willis Grimmia sp. unidentified Sarconeurum glaciale (C. Muell.) Card. & Bryhn

(Syn. Acarospora chlorophana (Wg.) Mass.)

(Syn. C. antarctica Filson)

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MARIE BYRD LAND ALGAE AND OTHER TERRESTRIAL BIOTA 223

490) suggested that a distinct band of Usnea high up the side of Washington Ridge, above lichen-free ground downslope, indicated recent ice recession, this and similar zonation on Mount Paterson appeared to be due to a geomorphological change from stable cliff-face and scree to less stable scree downslope.

Bryophytes

Mosses were generally sparse and infrequent. The species list (Table V) is preliminary. Previously, Baruam (1938) had found just two species from Washington Ridge. He found capsules on moss from Edsel Ford Range but none were seen during this survey. No liverworts have been found.

Mosses were not observed at Drummond Peak, Strider Rocks, Mount Schlossbach, La Gorce Peak, Clarke Peak and Mount Swadener despite careful searches, possibly due to absence of suitable moisture-retaining substrata. Elsewhere, most growths were small, scattered cushions no more than 10 cm across, usually on the moist mineral fines of frost- sorted ground. These were particularly frequent on the north-facing slopes of Breckinridge Peak. The largest confluent cushions, up to 30 cm wide on wet rocks, occurred on the knoll at the baseof Scott Nunataks and on inaccessible cliffs below nesting snow petrels on Mount Paterson.

Invertebrate fauna

Ciliate protozoa and testate amoebae were observed in several samples (Tables 11-IV). Testate amoebae, including Corythioa dubium Taranek, occurred amongst bryophytes and in lithosols with algal and lichen crusts. Vorticella sp. was found in a nutrient-enriched pond (sample 47) and unidentifiedciliates in lithosols. Three ciliates werecultured from bulked sample material; Leptopharym costatus Mermod, Pseudoplatyophrya nana (Kahl) Foissner and an unidentified hypotrich (possibly Urostyla sp.).

These appear to be the firstrecordsof Protozoa from Marie Byrd Land (Smith 1978, table 11) but are certainly not complete and a more detailed investigation is warranted. Elsewhere in Antarctica identical habitats support a more diverse protozoan fauna.

Rotifers and tardigrades were found (Tables 11-IV) but no nematodes, possibly due to extraction procedures not being used. Tardigrades occurred in bryophytes, lithosols and the nutrient-enriched pond at Mount Paterson. In addition to these habitats rotifers were encountered in the green crusts on stones and in cryoconite pools. Rotifers include Philodina gregaria and Philodina sp. in the two nutrient-enriched ponds (samples 46 and 47) and Adineta gracilis and Habrotrocha sp. in moist soils (samples 25, 29 and 33). Tardigrades include two species of Hypsibius (samples 25,

29,33 and47) and a single species of Echiniscus (samples 25 and 29).

No mites or collembola were found despite careful searching. Numerous mites were observed on similar green crusts at Cape Washington (Broady 1987, p. 12). Mites and collembola are widespread in continental Antarctica (Block 1984) as far south as 84'47's (Wise & Gressitt 1965) near Shackleton Glacier. However, they have not been found at Horlick Mountains andEllsworth Mountains (Wise &Gressitt 1965). The latter lie at a similar latitude to Edward VII Peninsula. Only a single species of mite is known in Marie Byrd Land (Strandtmann 1978,1981) from Ames andFloodranges. No difficulty was encountered there in finding this species below stones resting on moist sand. Environmental conditions must be very similar at Edward VII Peninsula which is only 3" further south. However, it lies about 600 km west of those records so perhaps dispersal barriers are operating. Under similar environmental conditions at Robenskollen, three mite species were found in high numbers on the under- surfaces of rocks but collembola were absent (Ryan & Watkins, in press).

Acknowledgements

This investigation would have been impossible without logistic support from Antarctic Division, DSKR, New Zealand and the US National Science Foundation. The help of my field companions, Peter Cleary, Chris Adam and Steven Weaver, is gratefully acknowledged. I thank T. Engelskjon and C. Howard-Williams for constructive criticisms of the manuscript.

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