Survey of Algae and Other Terrestrial Biota at Edward VII Peninsula, Marie Byrd Land

Antarctic Science 1 (3):2 15-224 (1989) 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, cryoconitepondscontained 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. 1 Reported here are observations on the algae, mosses, lichens Fig. 1. Location of Edward VII Peninsula and other regions and microfauna. A description of the ornithology has been mentioned in the text. 1-3 = Marie Byrd Land, 1 = Edward published elsewhere (Broady et a/. 1989). VII Peninsula, 2 = Edsel Ford Ranges, 3 = Ames and Flood Reports on the terrestrial biology of Edward VII Peninsula ranges, 4 = Horlick Mountains, 5 = Ellsworth Mountains, are few and lack detail. Early reports of mosses and lichens 6-8 = Dronning Maud Land, 6 = Robertskollen, were made by Presuud (in Amundsen 1912, p. 249), Gould 7 = Gjelsvikfjella and Miihlig-Hofmannfjella, (1931, p. 20) and Siple (1938, p. 490). The last author 8 = Unterseeoase, 9 =Liitzow-Holm Bay, 10 = Mawson Rock, 11 = Framnes Mountains, 12 = Vestfold Hills, reported mainly on more detailed studies made in Edsel Ford 13-14 = northern Victoria Land, 13 = Mount Kobourg, Ranges, 130 km further east (77"S, 145"W). Lichens were Birthday Ridge, 14 = Edmondson Point, Cape Washington, described by Dodge & Baker (1938) and mosses by Baruam Campbell Glacier, Inexpressible Island, 15 = southern (1938). Perkins(1945,p. 282) sampledplantsfrom 11 peaks Victoria Land dry valleys, 16 = McMurdo Ice Shelf, in Rockefeller Mountains. Lichens in these collections were 17 = Ross Island, 18 = Shackleton Glacier. 21 5

http://journals.cambridge.org Downloaded: 03 Mar 2015 IP address: 169.229.32.36 21 6 P.A. BROADY identified by Dodge (1973). Rudolph (1967) collected granites and metasediments. A general description of the lichens and noted an absence of microarthropods on region has been provided by Wade (1945) and a 1:250 000 Washington Ridge. Thisis theonly commenton microfauna map is available (United States Geological Survey 1972). although in eastern Marie Byrd Land, Siple (1938, p. 495) On 47 days in the field, during early to mid-summer, found protozoa, rotifers and tardigrades in pond water, and meteorological observations were taken twice a day at Strandtmann (1978,1981) described a new species of mite. approximately 1000and 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 Description of the region 76% of observations. Wind was frequent but did not exceed 20 kn. Conditions were calm for 13% of observations, The seven nunataks of the Alexandra Mountains and the 16 winds were <10 kn for 68% and >10 kn for 19%. The of theRockefeller Mountains are the only ice-free land in the prevailing wind was from east to north-east. Snowfall or Edward VII Peninsula, and comprise less than 0.5% of the snowdrift occurred on 25 days. 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 Methods and materials Nunataks are closest to the coast and are 11 km inland. Mount Paterson is furthest inland, 86 km from the coast. Exposed rock was reached at all nunataks except Mount Summits range from about 419 m to 1174 m altitude and, Manger where it was inaccessible. Efforts were made to except close to the coast, the surrounding ice-surfaces have collect the full diversity of vegetation at each nunatak. an elevation of 400 m to over 1000 m. The most extensive Samples of algae and substratum were collected into sterile nunataks, Mount Paterson and Washington Ridge in polythene bags using clean instruments. In the field, speci- Rockefeller Mountains (Figs 3-5), are low ridges up to 3 km mens were kept at ambient temperatures and were buried in long. Ice-free rock in Alexandra Mountains is much less snow at campsites. Field searches were made for mites and extensive and is mostly steep slopes and cliffs of two N-NE collembola using a lox hand-lens in likely habitats but no facing escarpments. The rocks of both ranges comprise 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

ROSS by Dr W. Foissner (Universiat Salzburg). Rotifers and

SEA t11.45 MtLBlRT ROCKS SlRIOtR ROCKS

MT PATERSON 77'30' EOWARO VII 0 +q-& .\MA 4.5.6.19.20.37 C'L 3 i PENINSULA \ Ill SCHLOSSBhCH i 22 50 PETREL 2.39 k23,38 4 46, FOKKLR ROCKS AE[KINRIDGt PK b ROCKEFELLER Ills \

Wr FRANKLIN' q5' km

906. WASHINGTON RIDGE Fig. 2. Geographical features of Edward VII Peninsula, Mane Byrd Land. Inset shows the location of Edward VII Peninsula 48,L9,4 in the south-east comer of Ross Sea. Nunataks of the MELT /d*30 Alexandra and Rockefeller mountains are shown as follows: N FLOW I 0 km 2 h SN = Scott Nunataks, SW = Mount Swadener, BO = Bowman L__ Peak, MA = Mount Manger, JO = Mount Josephine, CL = LAKE Clarke Peak, LG = La Gorce Peak, DR = Drummond Peak, FR = Mount Frazier, JA = Mount Jackling, FZ = Mount Fig. 3. Nunataks in the central Rockefeller Mountains, Mount Fitzsimmons, SH = Mount Shideler, PA = Mount Paterson Paterson and Washington Ridge are shown in Figs 4 and 5 (see Fig. 4), GO = Gould Peak, BU = Mount Butler, TE = respectively. Proglacial lakes and position of the Antarctic Tennant Peak. Nunataks to the east of Mount Paterson are petrel colony are indicated. Inset gives location (also see Fig. shown in more detail in Fig. 3. 2). Numbers refer to samples (see Tables i1-IV).

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

9% of occurrence and Algae abundance in all samples rare occ. freq. abund. TG - Cyanobacteria Aphanocapsa eiachista West & West 1 2 Chroococcus minor (Kuetz.) Naeg. 1 2 Chroococcus turgidus (Kuetz.) Naeg. 1 2 Cyanothece aeruginosa (Naeg.) Komarek 3 1 4 2 19 Gloeocapsa aipina Naeg. 2 1 18 Gioeocapsa kuetzingiana Naeg. 2 3 1 11 Gioeocapsa cf. punctata Naeg. 1 1 28 Gloeocapsa ralfsiana Kuetz. 2 1 2 1 11 Fig. 4. An aerial view of Mount Paterson taken from the north- cf. Chroococcidiopsis sp. 1 14 west at an altitude of about 1000 m. The nunatak is about Crinalium cf. magnum Fritsch & John 1 2 2.8 km lone. Oscillatoriaceae, trichomes 1-2 pm wide 2 16 Phormidium autumnale (Ag.) Gom. 2 2 1 9 Phormldium cf.fragiie (Menegh.) Gom. 1 2 Phormidium cf. laminosum Gom. 1 4 3 15 Schizothrir cf. antarctica Fritsch 12 Calothrir cf. parietina (Naeg.) Thuret 1 3 8 Homoeothrix cf. rivularis (Hansg.) Komarek & Kann 36 Nostoc sp. 1 1 3 5 19 Toiypothrix cf. bouteiilei (Breb. & Desm.) Forti 1 1 4 Stigonema minutum (Ag.) Hassall 1 26

Chlorophyta cf. Chioreiia SQ. 2 4 cf. Chlorococcum sp. 1 2 cf. Coenocystis sp. 1 2 1 3 13 cf. Dicfyochloropsis sp. 1 26 Pseudococcomyxa simplex (Mainx) Fott 4 5 17 view looking north along Washington Ridge from the Fig. 5. A Unidentified unicells 2 3 3 1 17 southernmost rock exposure. A frozen pond, which is Lichen gonidia with unicells 1 5 4 19 approximately 10 m long, can be seen in a windscoop in the Stichococcus baciilaris Naeg. 5 3 11 3 42 middle distance. cf. Desmococcus vulgaris Brand 2 3 5 14 45 Prasioia caiophyila (Carmich.) Menegh. 24 Prasiola crispa (Lightf.) Menegh. 1 5 11 tardigrades observed during microscopic examination of Prasioia uniseriate filaments 3 18 samples were identified by A. Suren (University of Canter- Prasiococcus caicarius (Boye Pet.) bury). Preliminary identificationsof lichens and bryophytes Vischer 4 3 13 were made by Dr R. Seppelt (Australian Antarctic Division) Chrysophyta and Dr R.I. Lewis Smith (British Antarctic Survey). cf. Botrydiopsis sp. 1 2

Results and discussion The flora, dominated by cyanobacteria and chlorophytes, is similar to that of the Vestfold Hills where 47% and 36% Algae of all species were cyanobacteria and chlorophytes respect- ively. Surprisingly, only a single chrysophyte was observed The algal flora of over 30 species (Table I) could probably and no diatoms were found as either living cells or empty be extended, especially for chlorophytes, by using cultures. frustules. This contrasts with other regions where diatoms Several of the present identificationsare tentative and require are usually found in both aquatic and non-aquatic habitats, cultures for confirmation. The algal flora of comparable e.g. ponds on Ross Island (Broady 1989, table 1) and ‘non- ‘terrestrialhabitats’ at Vestfold Hills has at least 77 species, aquatic’ habitats at Vestfold Hills (Broady 1986, table 6.5) of which 45 were found by microscopic examination of and nunataks (Pankow et al. 1987, p. 72, Ryan & Watkins, samples and the remainder from cultures (Broady 1986, in press). Although Engelskj@n(1986) does not mention dia- table 6.5). toms on nunataks in Dronning Maud Land they were present

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(O.M. Skulberg & T. Engelskjon, personal communication Epilirhic algae 1988). Three cyanobacteriaare of particular interest. Homoeothrix Thin black crusts of algae were observed infrequently where has been recorded only once before from the Antarctic melt-water trickled over the rock surfaces. In the Alexandra continent (Broady 1981a, p. 266, fig. 7). Specimens from Mountains black streaks down inaccessible cliff-faces were Edward VII Peninsula closely resemble H. rivularis KomArck seen at Mount Bowman, Mount Swadener and Scott Nunataks. & Kann 1973 (p. 206, figs 3847). There are only two Samples from Mount Franklin and Breckinridge Peak in previous records of Stigonema minutum (Broady 1986, p. Rockefeller Mountains (Table 11, samples 1 and 2) both 189, Ohtani & Kanda 1987,p. 261). Cyanothece aeruginosa contained abundant Gloeocapsa spp. The former was unusual has been recorded frequently from Antarctica (e.g. Broady in having as co-dominants Homoeothrix cf. rivularis and 1986, Pankow et al. 1987, Ohtani & Kanda 1987), however, Stigonema minutum. the occurrence of two size forms as suspected by Komkek Black epilithic crusts have a variable occurrence in coastal (1 976, p. 150) in material from Europe has not been confirmed continental Antarctica. They were not found at Edmondson before now. Measurements made on cells from sample 29 Point, Cape Washington and InexpressibleIsland in northern (Table 111) clearly demonstrated the occurrence of two Victoria Land (Broady 1987a) but did occur at Vestfold populations centred around cell widths of about 10 pm and Hills and Mawson Rock (Broady 1981~).At these locations and 16 pm. and in Dronning Maud Land (Engelskjon 1986, Ryan & In the following sections algal communities are described Watkins, in press) Gloeocapsa spp. were dominant as at from the full range of habitats at Edward VII Peninsula and Rockefeller Mountains. However, Homoeothrix sp. has are compared with those elsewhere in Antarctica. Algae in been recorded previously only at Mawson Rock (Broady individual samples are listed in Tables 11-IV and locations of 198 1a). samples are shown in Figs 2 and 3. Although absent at Alexandra Mountains, at Rockefeller

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

Sample material, locatlon and number Epilithic Chasmoendolithic Algae Rockefeller Mountains Rockefeller Mountains Alexandra Mountains FA' BR DR NI NI NI NI FA DR DR ST PA JA FR LG SW SN SN 1 2 345618 9 10 11 12 13 14 15 16 17 18 - Cyanobacteria Cyanothece aeruginosa Gloeocapsa alpina a 0 Gloeocapsa kuetzingiana a f Cioeocapsa cf. punctata 0 a Gioeocapsa ralfsiana a r cf. Chroococcidiopsis a Oscillatoriaceae, trichomes 1-2 pm wide Phormidiwn auiumnale Phormldium cf. laminosum f Calothrix cf. parietina 0 0 Homoeothrix cf. rivularis a Nostoc sp. af Sfigonema minutum a

Chlorophyta cf. Coenocystis sp. 0 f a Pseudococcomym simplex a Unidentified unicells 0 ff a Lichen gonidia with unicells f affa a f Stichococcus bacillaris fff ff+ ro cf. Desmococcus vulgaris f a aaffa aa

Microfauna Rotifers + ++

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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Mountains green algal crusts and associated lichens on rock types or to lower moisture availability in that high stones on scree slopes were much more frequent than black altitude desert. 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 Soil algae Ridge. They also occurred where melt-water percolated downslope, mostly on the shaded and relatively moist lowcr Soils were either raw mineral soils (lithosols), or nutrient- sides and undersurfaces of the superficial layer of stones. enriched (ornithogenic) in which there was a substantial All six samples (Table 11, samples 3-8) were either domi- organic component derived from bird guano, feathers and nated by lichens or had lichens as a major component. The carcasses. Lithosols were very restricted in extent and lichens had enhanced development of the unicellular green occurred as small areas of mineral fines on sorted ground, as phycobiont and superficially appeared to be green crusts of deposits in crevices and on ledges, and as coarse gravelly free-living algae. Associated free-living algae were products of decompositionof the more friable granitic rocks. predominantly Stichococcus bacillaris, cf. Desmococcus Ornithogenic soils occurred in the vicinity of Antarctic vulgaris and Pseudococcomyxa simplex. petrel and snow petrel nests on Washington Ridge and Similar green algal crusts have been described at Cape Mount Paterson in Rockefeller Mountains and amongst Washington, northern Victoria Land (Broady 1987a, p. 12) abandoned nesting sites (Broady et at. 1989) on the lowest and on Ross Island (Broady 1989). However, at both those rocky knoll of Scott Nunataks. coastal locations the crusts were dominated by free-living Lichens and algae were occasionally visible as thin black chaetophoralean algae, including specimens resembling cf. crusts over lithosols. The most frequent algal components D.vulgaris of this study. (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. Chasmoendolithic algae Coenocyslis sp. amongst gravel and small stones were encountered twice (Table 111, samples 19 and 28). Water- A striking feature at granitic nunataks was the abundance of flushed soils were rare but on Mount Fitzsimmons and chasmoendolithic lichens and less frequently of free-living Mount Jackling narrow gullies containing wet gravel supported algae (Table 11, samples 9-18). The former grew in narrow ribbon-like thalli of Prasiola calophylla (Table 111, samples cracks between and below translucent crystals of quartzite 26 and 27). and feldspar. They were mostly absent from opaque Foliaceous Prasiola crispa occurred downslope from the metasediments except where these contained veins of quartzite large colony of Antarctic petrels on the southernmost out- as on Drummond Peak and La Gorce Peak. The free-living crop of Mount Paterson (Fig. 3; Table 111, samples 31 and algae usually formed green crusts and cf. D. vulgaris was 32). Mats covered approximately 500-1000 m2 where the often dominant. Cyanobacteria were infrequent but Nostoc ground received melt from adjacent snow drifts. The only sp., Gloeocapsa cf. punctata and cf. Chroococcidiopsis sp. other occurrence of P. crispa was at Scott Nunataks (Table were each abundant in a single sample. 111, samples 33, 35 and 36) near the abandoned nest sites. Endolithic algae were scarce at Robertskollen due to Frequent microscopic associates were cf. D. vulgaris and P. unsuitable rock types (Ryan & Watkins, in press) however, calcarius. Also, these two species dominated the green ‘Pleurococcus’ was frequent at more easterly nunataks crusts on a gravel soil close to the nesting sites of snow (Engelskjfln 1986, p. 217). This group includes species petrels on Washington Ridge (Table 111, sample 30). similar to cf. D. vulgaris and to Desmococcus sp. A, a Algae populations on lithosols were small in comparison chasmoendolith at Mawson Rock and nearby nunataks in with coastal regions of Antarctica, e.g. Vestfold Hills (Broady North Masson Range and at Vestfold Hills (Broady 19816, 1986,~.18l)andRossIsland (Broady 1989) wheresaturated p. 263, table 1). At coastal Mawson Rock, Desmococcus sp. ground is often covered with mats of Oscillatoriaceae and A occurred only in areas not swept by sea-spray. In areas mucilaginous colonies of Nostoc. The cause could be the receiving salineaerosolsP. calcarius was dominant (Broady absence of poorly drained substrata as most slopes were 1981b, table 2). The generally low salt environment at covered with boulders and well-drained stoney ground. At inland nunataks appears ideal for the growth of cf. D. Gjelsvikfjella and Miihlig-Hofmannfjella there was more vulgaris and similar species. However, the dominant endo- extensive wet ground with mats of these cyanobacteria lithic algae at these locations are in contrast to those at (Engelskjfln 1986, table 2, p. 217) . upland sites with similarly low salt levels in southern Victoria No other study has recorded such a high frequency of C. Land where a unicellular chlorophyte, Hemichloris antarctica aeruginosa in lithosols despite its occurrence in several Tschermak-Woess & Friedmann, is often accompanied by regions, both coastal (Broady 1986,table 6.5,1987a, table 1, coccoid cyanobacteria (Friedmann & Ocampo-Friedmann Ohtani & Kanda 1987, p. 259) and at inland nunataks 1984, p. 180). This difference could be related to differing (Pankow et al. 1987, p. 69, Ryan & Watkins, in press). In

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

Sample material, location and number Algae Lithosols Bird-influenced '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

Cyanobacteria Chroococcus minor r Cyanothece aeruginosa o ffra a Gloeocapsa kuetzingiana f 0 f Gloeocapsa cf. punctata r Gloeocapsa ralfsiana f Oscillatoriaceae, trichomes 1-2 um wide a Phormidium auumnale 0 r f Phormrdium cf. laminosum f a Schizothrir cf. antarctica a Calothrix cf. parietina 0 Nostoc sp. r a Stigonema minutum ar

Chlorophyta cf. Chiorella sp. f cf. Coenocystis sp. a a 0 Pseudococcomyxa simplex 0 0 a 0 Unidentified unicells ro r Stichococcus bacillaris f of r r a cf. Desmococcus vulgark or a aa aa f aa Prasiola calophylla aa Prasiola crispa aaa aa Prasiococcus calcarius . fa aff

Chrysophyta Botrydiopsis sp 0

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

' 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 and on Ross Island (Broady 1989). common, especiallyHantzschia amphioxys (Ehr.) Grun. and Navicula muticopsisvan Heurck,e.g. Vestfold Hills (Broady 1986, table 6.4), and southern Victoria Land (Seaburg et al. Algae epiphytic on moss 1979, table 2). N. muticopsis can also be abundant amongst P.crispa on bird-influenced soils, e.g. Vestfold Hills (Broady Moss cushions were frequently covered with black crusts of 1986, p. 183) and Inexpressible Island (Broady 1987a, p. lichens and algae. The epiphytic algal flora (Table IV, 13), and its absence at the Mount Paterson bird colony is samples 37-44) was dominated by Nostoc sp., C. aeruginosa unusual. P. crispa iscommon near birdcolonieson nunataks and P. simplex. Moss cushions invariably bear crusts of (Engelskjen 1986, p. 218, Ryan & Watkins, in press). epiphytic algae and the flora recorded here is not unusual The occurrence of P. calcarius on bird-influenced soils is (cf. Ohtani & Kanda 1987) except for the absencc of dia- probably related to high salt levels rather than to nutrient- toms. Although rarely dominant, diatoms are usually fre- enrichment. The preference of this alga for saline environments quent, especially Pinnufaria borealis Ehr. (Broady 1982, p. without bird-influence has been noted in Europe (Lund 222, 1986, p. table 6.4, Ohtani 1986, p. 218). P. simplex 1966, p. 307) and Antarctica (Broady 1983, p. 216). More appears to favour moss as a substratum at Edward VII recently it has been found on ground close to penguin Peninsulaas it does elsewhere in Antarctica (Broady 1987b, rookeries in northern Victoria Land (Broady 1987a, p. 13) table 1).

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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 Algae Bryophytes Ponds Other NI' FO BR BU SN SN SN SN 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

Cyanobacteria Aphanocapsa elachista 0 Chroococcus turgidus r Cyanothece aeruginosa f 1 Gloeocapsa alpina r Gloeocapsa kuetzingiana 0 Gloeocapsa cf. punctata a Gloeocapsa ra[fsiana 0 f Crinaliwn cf. magnum r Oscillatoriaceae,trichomes 1-2 pm wide Phormidium autumnale Phormidium cf. fragile 0 Phormldiwn cf. laminosum a af of Calothrix cf. parietina Homoeothrix cf. rivularis aa NOSfOC sp. 0 f a f Tolypothrix cf. bouteillei f

Chlorophyta cf. Chlorella sp. f cf. Chlorococcum sp. cf. Coenocystis sp. r cf. Dictyochloropsis sp. aaf Pseudococcomyxa simplex a a a Unidentified unicells f Lichen gonidia with unicells o f Stichococcus bacillaris 0 f af ffa cf. Desmococcw vulgaris fr ao Prasiola crispa f Prasiola crispa uniseriate filaments a Prasiococcus calcarius af

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

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

Pond algae Another much larger nutrient-enrichedpondoccurred on ice at the base of the Antarctic petrel rookery on Mount Paterson Aquatic habitats were rarely encountered. Small but com- (Fig. 3, Table IV, sample 47) and contained abundant pletely frozen ponds in windscoops were seen on Mount uniseriate P. crispa and felts of P. cf. faminosum. The algal Paterson, Washington Ridge and Mount Jackling but were floraofthesenutrient-enrichedponds is not unusual although absent from Alexandra Mountains. Other than these, only it differs from the dense populations of Chlamydomonas sp. two ponds were encountered on ice-free ground, both in in a petrel rookery pool in Muhlig-Hofmannfjella(Engelskjon Rockefeller Mountains (Table IV, samples 45 and 46). One 1986, p. 217) and in penguin rookery pools on Ross Island was a 10 cm deep pool in a granite boulder at Strider Rocks (Broady 1989, table 3). in which the only algae were Gloeocapsa ralfsiana and S. The most extensive areas of open water were two shallow bacillaris. The other was a nutrient-enriched pool close to lakes in ice basins 1.5 and 0.5 km south-west of Washington nesting skuas on the middle outcrop of Fokker Rocks. This Ridge and Mount Paterson respectively (Fig. 3). At the time contained rich red-brown felts of Phormidium cf. laminosum. of our visit to Washington Ridge, substantial melt-water was

http://journals.cambridge.org Downloaded: 03 Mar 2015 IP address: 169.229.32.36 222 P.A. BROADY pouring into the lake, c. 300 m wide, from a wide area of in abundance only where they would receive moisture from ablating ice which had received wind-blown sand and gravel melting snowdrifts, for instance on ledges, in depressions blown off the nunatak. The cryoconite ponds in this area and amongst boulders on scree slopes e.g. at Drummond were the most extensive freshwater habitat containing visible Peak where numerous small snowdrifts were scattered over algae (Table IV, samples 48-50). Abundant, small, dark the boulder slopes and were the only water source for the brown wefts of H. cf. rivularis, cf. Dictyochforopsis sp. and abundant epilithic lichens. Lithosols also had black crusts S. bacillaris filaments occurred amongst the mineral material. of lichens, especially where the soils were in slight de- Identical, although less extensive pools were found south- pressions attracting drift snow. west of Mount Paterson adjacent to a line of moraine. The Abundant chasmoendolithic lichens were present in coarse- flora was similar except for the absence of H. cf. rivularis grained granites containing large feldspar crystals, e.g. at and the presence of Nostoc sp. Mount Frazier, Mount Jackling, Mount Fitzsimmons and Homoeothrix has not been observed previously in cryoconite Mount Shideler in northem Rockefeller Mountains. Epilithic pols, although Nostoc and Oscillatoriaceae are common in lichens were often sparse on these more rapidly weathering other regions (Wharton et al. 1981, Broady 1987a, table 1). granites whereas richer growths occurred on more stable The absence of diatoms is unusual as these are often abundant micro-granites and in particular on schists, e.g. on Mount elsewhere. Pinnularia cymatopleura West & West is Nilsen, Breckinridge Peak, Drummond Peak, Clarke Peak particularly characteristic of this habitat, e.g. on McMurdo and Scott Nunataks. Ice Shelf (Kellogg & Kellogg 1987, p. 85), Ross Island Abundant growths of large lichen thalli were present (Broady 1989, table 1) and on Campbell Glacier, northern immediately adjacent to the ice edge at Mount Frazier, Victoria Land (Broady 1987a, p. 15). 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. Algae porn other habitats

Table V. A preliminary list of lichens and bryophytes from Edward VII Occasionally bird bones were found supporting green algal Peninsula, Marie Byrd Land crusts dominated by S. bacillaris, cf. D. vulgaris and P. calcarius (Table IV, samples 52 and 53). The latter two Lichens algae were common in this niche at Vestfold Hills (Broady Biatorellu cerebriformk Filson 1986, p. 184, figs 28, 29). P. calcarius also formed rich (Syn. Acarospora chlorophana (Wg.) Mass.) Buellia frigido Darb. crusts over the surface of an old cache of dog food left by the cf. Bueliia soredians Filson Second Byrd Expedition of 1933-35 at the northern end of Buellia sp. Mount Franklin (Table IV, sample 41). Caloplaca athallina Darb. Caloplaca sp. Candelariella hallettensis (Murray) Ovst. Lichens (Syn. C. antarctica Filson) Lecidea physciellu Darb. Lecidea sp. All nunataks had lichens; overall they were the most abundant cf. Lecanora sp. vegetation. Species listed (Table V) represent a preliminary cf. Lepraria sp. examination of only a small proportion of the large number Physcia cuesia (Hoffm.) Hampe Pseudephebe minuscula (Nyl. ex Amold) Brodo & Hawksw. of specimens, which clearly contain other as yet unidentified Pseudephebe aff. pubescens 6.)Brodo & Hawksw. species. The most prominent species were Pseudephebe Rhizocarponflavum Dodge & Baker minuscula, Umbilicaria decussata and Usnea sphacelata, Rhizocarponflavum f. subfoliosum Filson with Caloplaca spp. and Xanthoria spp. near nesting birds. Rhizoplaca melanophthalma (Ram.) Lwck. & Poelt Thedominant speciesresemblethoseof GroupEdescribed Rinodinu olivaceobrunnea Dodge & Baker Umbilicaria aprina Nyl. by Pickard & Seppelt (1984, table 3, p. 94). This community Umbilicariu decursata (Vill.) Zahlbr. grows close to the polar ice sheet where there is relatively Usnea sphacelata R. Br. high moistureavailabilityand little wind-blown salt. Similar Xanthoria elegans (Link.) Th. Fr. communities were described by Kappen (1985) in northern Xanthoria muwsonii Dodge Victoria Land although Usnea was very rare on Mount Mosses Kobourg, the inland nunatak, but abundant at coastal Birthday Bryum argenteum Hedw. Ridge. Bryum pseudotriquetrum (Hedw.) Schwaegr. Excellent descriptive observations on the ecology of lichens Grimmia antarcfici Card. in Marie Byrd Land have been made by Siple (1938) and Grimmia lawiana Willis little can be added to these. Locally, water availability was Grimmia sp. unidentified Sarconeurum glaciale (C. Muell.) Card. & Bryhn a major factor limiting lichen growth. Epilithic lichens grew

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490) suggested that a distinct band of Usnea high up the side 29,33 and47) and a single species of Echiniscus (samples 25 of Washington Ridge, above lichen-free ground downslope, and 29). indicated recent ice recession, this and similar zonation on No mites or collembola were found despite careful searching. Mount Paterson appeared to be due to a geomorphological Numerous mites were observed on similar green crusts at change from stable cliff-face and scree to less stable scree Cape Washington (Broady 1987, p. 12). Mites and collembola downslope. 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 Bryophytes Mountains andEllsworth Mountains (Wise &Gressitt 1965). The latter lie at a similar latitude to Edward VII Peninsula. Mosses were generally sparse and infrequent. The species Only a single species of mite is known in Marie Byrd Land list (Table V) is preliminary. Previously, Baruam (1938) (Strandtmann 1978,1981)from Ames andFloodranges. No had found just two species from Washington Ridge. He difficulty was encountered there in finding this species found capsules on moss from Edsel Ford Range but none below stones resting on moist sand. Environmental conditions were seen during this survey. No liverworts have been must be very similar at Edward VII Peninsula which is only found. 3" further south. However, it lies about 600 km west of those Mosses were not observed at Drummond Peak, Strider records so perhaps dispersal barriers are operating. Under Rocks, Mount Schlossbach,La Gorce Peak, Clarke Peak and similar environmental conditions at Robenskollen, three Mount Swadener despite careful searches, possibly due to mite species were found in high numbers on the under- absenceof suitable moisture-retainingsubstrata. Elsewhere, surfaces of rocks but collembola were absent (Ryan & most growths were small, scattered cushions no more than Watkins, in press). 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 Acknowledgements confluent cushions, up to 30 cm wide on wet rocks, occurred on the knoll at the baseof Scott Nunataks and on inaccessible This investigation would have been impossible without cliffs below nesting snow petrels on Mount Paterson. 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 Invertebrate fauna Weaver, is gratefully acknowledged. I thank T. Engelskjon and C. Howard-Williams for constructive criticisms of the Ciliate protozoa and testate amoebae were observed in manuscript. several samples (Tables 11-IV). Testate amoebae, including Corythioa dubium Taranek, occurred amongst bryophytes and in lithosols with algal and lichen crusts. Vorticella sp. References was found in a nutrient-enriched pond (sample 47) and unidentifiedciliates in lithosols. Three ciliates werecultured AMIJNDSEN,R. 1912. The South Pole, London: John Murray, 449 pp. BARTRAM,E.B. 1938. Botany of Second Byrd Antarctic Expedition 111. from bulked sample material; Leptopharym costatus Mermod, Mosses. Annals ofthe Missouri Bofanic Garden, 25,719-724. Pseudoplatyophrya nana (Kahl) Foissner and an unidentified BLOCK,W. 1984. Terrestrial microbiology, invertebrates and ecosystems. In hypotrich (possibly Urostyla sp.). LAws,R.M.,ed.Anfarcticecology, 1.London: Academic Press, 163-236. These appear to be the firstrecordsof Protozoa from Marie BROADY,P.A. 1981a. Ecological and taxonomic observations on subaerial Byrd Land (Smith 1978, table 11) but are certainly not epilithic algae from Princess Elizabeth Land and Mac.Robertson Land, complete and a more detailed investigation is warranted. Antarctica. British Phycological Journal, 16,257-266. BROADY,P.A. 19816. The ecology of chasmolithic algae at coastal locations Elsewhere in Antarctica identical habitats support a more of Antarctica. Phycologiu,20,259-272. diverse protozoan fauna. BROADY,P.A. 1982. Ecology of non-marine algae at Mawson Rock, Rotifers and tardigrades were found (Tables 11-IV) but no Antarctica. Nova Hedwigia, 36,209-229. nematodes, possibly due to extraction procedures not being BROADY,P.A. 1983. The Antarctic distribution and ecology of the terrestrial used. Tardigrades occurred in bryophytes, lithosols and the chlorophytan alga Prasiococcus calcarius (Boye Petersen) Vischer. Polar Biology, 1.21 1-216. nutrient-enriched pond at Mount Paterson. In addition to BROADY,P.A. 1986. Ecology and taxonomy of the terrestrial algae. In these habitats rotifers were encountered in the green crusts RCKARD,J., ed. Anfarcficaasb. Sydney: Academic Press, 165-202. on stones and in cryoconitepools. Rotifers includePhilodina BROADY,P.A. 1987a.A floristic survey of algae at four locations in northern gregaria and Philodina sp. in the two nutrient-enriched Victoria Land. New Zealand Antarctic Record, 7 (3). 8-19. BROADY,P.A. 19876. The morphology, distribution and ecology of ponds (samples 46 and 47) and Adineta gracilis and Pseudococcomyxa simplex (Mainx) Fott (Chlorophyta, Chlorellaceae), a Habrotrocha sp. in moist soils (samples 25, 29 and 33). widespread terrestrial Antarctic alga. Polar Biology, 7,25-30. Tardigrades include two species of Hypsibius (samples 25, BROADY,P.A. 1989. Broadscale patterns in the distribution of aquatic and

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terrestrial vegetation at three ice-free regions on Ross Island, Antarctica. 209-219. Hydrobiologia, 172,77-95. OHTANI,S. & KANDA,H. 1987. Epiphytic algae on the moss community of BROADY,P.A., ADAMS,C.J., CLEARY,P.J. & WEAVER,S.D. 1989. Grimmia lawiana around Syowa Station, Antarctica. Proceedings of the Ornithological observations at Edward VII Peninsula, Antarctica, in Nafional Institute of Polar Research Symposium on Polar Biology, So. I 1987-88. Notornis, 36,53-61. 255-264. DODGE,C.W. 1973. Lichen flora of the Antarctic continent and adjacent PANKOW,€1. VON, HAENDAL,D.,RIcHER, W. & WAND,U. 1987. Algologische islands. Canaan: Phoenix, 399 pp. Beobachtungen in der Schirmacher- und Lnterseeoase (Dronning Maud DODGE,C.W. & BAKER,G.E. 1938. Botany of Second Byrd Antarctic Land, Ostantarktika). Archiv fiir Protistenkunde, 134,59-82. Expedition 111. Lichens and lichen parasites. Annals of the Missouri PERKINS,J.E. 1945. Biology at Little America 111, the West Base of the United Botanic Garden, 25,515-718. States Antarctic Sew ice Expedition 1939-41. Proceedings of the ENGELSKJ0N, T. 1986. Botany of two Antarctic mountain ranges: American Philosophical Society, 89,270-284. Gjelsvikfjella and Miihlig-Hofmannfjella, Dronning Maud Land. I. RCKARD,J. 1986. Antarctic oasis. Sydney: Academic Press, 367 pp. General ecology and development of the Antarctic cold desertcryptogam Pic~am,J. & ADAMSON,D.A. 1983. Perennially frozen lakes at glacierhock formation. Polar Research, 4,205-224. margins, East Antarctica. In OLIVER,R.L., JAMS, P.R. & JAGO,J.B., eds. FRIEDMANN,E.I. 1982. Endolithic microorganisms in the Antarctic cold Antarcric earfh science. Canberra: Australian Academy of Science & desert. Science, 215, 1045-1053. Cambridge: Cambridge University Press, 470-472. FREDMANN,E.I. & OCAMPO-FREDMANN,R. 1984. Endolithicmicroorganisms RCKARD,J. & SEPPELT,R.D. 1984. Phytogeography ofAntarctica. Journal o/ in extreme dry environments - analysis of a lithobiontic microbial Biogeography, 11.83-102. community. In KLUG,M.J. & REDDY,C.A., eds. Current perspectives in RUDOLPH,E.D. 1967. Biological survey of Marie Ryrd Land. Antarctic microbial ecology. Washington,DC: American Society for Microbiology, Journal of the United Stales, 2 (4),96-97. 177-185 pp. RYAN,P.G. & WATKINS,B.P. In press. Biological survey of Robertskollen, GEITLER, L. 1932. Cyanophyceae. Kryptogamenjlora, 14. Liepzig: westem Dronning Maud Land: area description and preliminary species Akademische Vorlagsgellschaft, 1196 pp. lists. South African Journal of Antarctic Research. GOULD,L.M. 1931. Cold, the record of an Antarctic sledge journey. New SEABURG,K.G.,PARKEK, R.C., PREscon, G.W. & WHITFORD,L.A. 1979. The York: Brewer, Warren & Putman, 275 pp. algae of southern Victoria Land, Antarctica. A taxonomic and HERMICHEN,W.-D., KOWSKI,P. & WAND,U. 1985. Lake Untersee, a first distributional study. Bibliothecia Phycologica, 46, 169 pp. isotope study of the largest freshwater lake in the interior of East SIPLE,P.A. 1938. The Second Byrd Antarctic Expedition - botany. Annals Antarctica. Nature, 315, 131-133. of the Missouri Botanic Garden, 25,467-5 14. KAPPEN,L. 1985. Vegetation and ecology of ice-free areas of northern SMTH,H.G. 1978. Thedistributionand ecology ofterrestrialProtozoaof sub- Victoria Land, Antarctica 1. The lichen vegetation of Birthday Ridge and Antarctic and maritime Antarctic islands. British Antarctic Survey an inland mountain. Polar Biology, 4,213-225. Scientific Reports, No. 95, 104 pp. KEI.I.OOG,D.E. & GLLOGG,T.B. 1987. Diatoms of the McMurdo Ice Shelf STRANDTMANN,R.W. 1978. Terrestrial arthropods, Marie Byrd Land, implications for sediment and biotic reworking. Palaeogeography, Antarctica. Antarctic Journal ofthe United States, 13 (4), 166-168. Palaeoclimatoiogy, Palaeoecology, 60,77-96. S~RANDTMANN,R.W. 1981. Kotes on Nanorchestes I. Description of KOMAREK,J. 1976. Taxonomic review of the genera Synechocystis Sauv. Nunorchestes wifbanksi n. sp. (Acari: Nanorchestidac) from Marie Byrd 1892, Synechococcus Naeg. 1849, and Cyanothece gen. nov. Land, Antarctica. Pacific Insects, 23, 107-1 11. (Cyanophyceae). Archiv fiir Protistenkunde, 118, 119-179. UNITED STATESGEOLOGICAL SURVEY. 1972. Alexandra Mountains,Antarctica. KOMAREK,J. & KANN,E. 1973. Zur Taxonomie und C)kologie der Gattung 1:250 000. Washington DC: United States Department of the Interior, Homoeothrix. Archiv fiir Protirtenkunde, 115,173-233. Geological Survey. LEND,J.W.G. 1966. A genus new to Poland: Prasiococcus Vischer. Acta WADE,F.A. 1945. The geology of the Rockefeller Mountains, King Edward Hydro biolog ia, 8,303-3 09. VII Land, Antarctica. Proceedings of the American PhilosophicalSociety, @UNI, A., TANAKA,Y., UEMURA, M., YAMAMOTO,M. & TAKAHASHI, E. 1987. 98.67-77. Floristic studies on algaefrom inland waters of Ongul Islandsand vicinity, WHARTON,R.A. JR, VINYARD, W.C., PARKER,R.C., SIMMOSS,G.M. JR. & Antarctica: I. East Ongul Island. Proceedings ofthe National Institute oj SEABURG,K.G. 1981. Algae in cryoconite holes on Canada Glacier in Polar Research Symposiwn on Polar Biology, No. I, 233-254. Southern Victorialand, Antarctica. Phycologia, 20,208-21 1. OHTANI,S. 1986. Epiphytic algae on mosses in the vicinity of Syowa Station. WISE,K.A.J. & GRESSIIT,J.L. 1965. Far southem animals and plants. Nature, Memoirs oft he Nationallnrtitute offolar Research, Speciallssue, No. 44, 207, 101-102.

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