Tardigrada from the Husvik area, South Georgia, subAntarctic

PREBEN S . OTTESEN AND TERJE MElER

Ottesen, P. S. & Meier, T. 1990: Tardigrada from the FIusvik arca, South Georgia, sub-Antarctic. Polar Research 8 , 291-294. Eleven species of tardigrades in South Georgia, of which two are new to science, were found in samples collected at fifteen localities. The highest number of species was found in moss from a scree field. Twenty species of tardigrades arc presently known from South Georgia, but the island remains insufficiently investigated. The species composition is similar to that of southern South America. The high number of cosmopolitan species makes the geographical distribution pattern of the South Georgian tardigrades more similar to that of macrolichens than to that of insects and vascular plants.
Preben S . Ottesen, Universiry of Oslo, Department of Biology, Division of Zoology, P . O . Box 1050 Blindern, N-0316 Oslo 3, Norway, present address: The Agricultural University of Norway, Department of Biology and Nature Conseroation, P. 0 . Box 14, N-1432, A S - N L H , Norway; Terje Meier, Lensmann Hiorths all6 3 , N-0661 Oslo 6 , Norway; October 1989 (revised February 1990).

While the tardigrades of the continental and maritime Antarctic have been extensively studied (i.e. Murray 1906, 1910; Jennings 1976a, b; Everitt 1981; Dastych 1984; McInnes & Ellis-Evans 1987; Usher & Dastych 1987; Miller et al. 1988), few studies have been carried out in the sub-Antarctic. Richters (1920), Iharos (1963) and MihelEit (1967) identified tardigrades from the southern part of South America, Richters (1920) from the Falkland Islands and Richters (1920), Jennings (1976b) and Dastych (1984) from South Georgia. Previous sampling at South Georgia has been done in the Royal Bay and Cumberland Bay areas, particularly at King Edward Point, and in Grytviken. This paper reports species found by the British Antarctic Survey terrestrial ecology expedition to Husvik in the Stromness Bay area on South Georgia in 1988 (Block et al. 1988).

The samples were extracted according to the method of Morgan & King (1976), i.e. soaked in water for 24 hours, then squeezed dry over a Petri dish, resoaked and allowed to stand for several minutes, then squeezed again. The tardigrades were removed from the water in the Petri dish with a capillary tube under 30 x magnification and transferred to a microscope slide. They were mounted in Hoyer's medium and identified using a compound microscope at 400 and 1,000 x magnifications. The primary identification manual was Ramazzotti & Maucci (1983). Terje Meier identified the tardigrades and supplied information on distribution and ecology.

Results
A description of the fifteen localities, from the wettest to the driest communities, is given in Table 1. Their distribution in the Husvik area is shown in Fig. 1. A list of the tardigrades in the various samples is given in Table 2. Approximately half of the samples, among them the dense moss banks which form deep peat deposits (loc. 10 and 11), contained no tardigrades. We had hoped to find some tardigrades living in gravel and sandy soil (loc. 12 and 13) as this habitat has been reported recently to contain interesting species in the alpine grassland of Italy (Manicardi & Bertolani 1987) and on Dundee Island in the maritime Antarctic (Usher & Dastych 1987). In

Material and methods

The samples for tardigrade extraction were collected between 11 and 13March 1988 northwest of the abandoned Husvik whaling station (54"10'S, 36'42'W) and at the Tonsberg Point peninsula at altitudes from sea level to 8 0 m a.s.1. (Fig. 1). Each sample had a volume of approximately 300cm3. Each was placed in a plastic bag and kept in a refrigerator at 4C during transport to Rio d e Janeiro, arriving in Norway one day later.

292 P . S. Ottesen & T. Meier

Table 1. List of sampling localities, from the approximately wettest (1) to the driest (15) habitat.
I : Pond, 1 0 x 4 m . 1 m deep, 30m a.s.1.. surrounded by Rostkooiu vegetation at both ends, gravel and stones on sides. Bottom with mosses and algae, some spots with sand. Thc sample contained mud from the bottom. 2: As in 1 , but the sample consisted of aquatic mosses. 3: Moss-sample from bog dominated by Juncus scheuchrerioides, i t . a wet nutrient rich bog. 4: Moss-sample from bog with J . scheuchzerioides, Rostkooia rnagellanica and herbs; drier than 3. 5 : Moss growing among and above stones along a small creek. 6: Moss growing among stones close to the water linc at Reservoir Lake (old water reservoir for the Husvik whaling station). 7: Moss growing below the dam at Reservoir Lake, sprayed with water from the outlet waterfall. 8: Steep cliff, exposed towards the south, water dripping down the wall. Partly covered with a rich vegetation of mosses and lichcns, several grasses and herbs, such as Acaena. The sample consisted of mosses. 9: As in 8, but sample consisted of lichens. 10: Moss hank, c . 10 m a . s . l . ,with dense stands of mosses of thc genera Polyrrichum and Dicrunum. covzringlarge areas. 11: As in 10, but c. 50w.a.s.l. 12: Soil bample from grassland dominated by Festuca contracla. Dry soil. 13: As in 12. 14: Scree-field with some mosses, Acuena and scattered grasses at c. 30 m a.s.1. Sample contained rnosscs. 15: Temporary pond, at 80m a.5.l. Dry whcn samplcd. Sample consisted of mosses.

only one of our sandy soil samples (loc. 12) was a tardigrade found, Macrobiotus furciger, which is a common habitat generalist. Localities 2, 4, 5, 6 and 7 contained no tardigrades, perhaps due to the sampling procedure, or other unexplained factors. LOC.14, moss from a scree field, proved to be the richest habitat in terms of number of species and individuals. One of these species, Isohypsibius sp., is new to science. It resembles I. improvisus, but has different claws. Another species new to science was found in moss of a dry pond (loc. 15). This Macrobiotus sp. resembles M . blocki, but the eggs are different (R. Mobjerg Kristensen pers. comm.). Both of these new species will be described by Terje Meier in a separate publication.

Discussion
Table 3 provides a survey of the tardigrades known from South Georgia. Prior to the present study fifteen species were known, of which we found six, while five species are now reported 21s new to the island. Currently, twenty species of tardigrades are known from South Georgia. There is no doubt that more species of tardigrades exist on South Georgia. Despite the scanty material. some patterns of the colonization of the South Georgia tardigrade fauna begin to appear. Macrobiotus hufelandi and Milnesiurn tardigradum are true cosmopolitan species, while M . furciger, Hypsibius dujardini, Ramazzottius oberhaeuseri and Diphascon pingue are fairly widespread throughout the world (Ramazzotti 6 Maucci 1983), including previous ( records from continental or maritime Antarctic. Macrobiotus liiiiae, Hypsibius pullidus and Isohypsibius prosostomus also have a world wide distribution, but they have not been reported from the Antarctic. Mopsechiniscus imberbis is a South American species. Although South Georgia has been poorly investigated, it is striking that a number of very common and widespread Antarctic species mentioned by Dastych (1984) such as Echiniscus pseudowendti, E . jwningsi, Hypsibius renuudi and Diphascon puniceus, have not yet been found there. Although it is dangerous to base conclusions on negative records, it would seem likely that the original distribution centre of the South Georgian tardigrade fauna was to be sought in the Falkland Islands and South America

.D

Fig. 1. Map of the sampling arca. Locality numbers refer to Table 1. Shaded areas are lakes or ponds. Altitude in mctrcs.

Tardigrada from the Husvik area, South Georgia, sub-Antarctic 293

Table 2. List of tardigrade species from Husvik in systematic order. Locality numbers refer to Table 1. The numbers in the table refer to the number of individuals found, while X denotes common, XX very common and - absent.

Farnily/Species
1 2 3 4 5

Locality
6
7 8 9 1 0 1 1 1 2 1 3 1 4 1 S
~~ ~

Echiniscidae Mopsechiniscus imherhis (Richters, 1920) Hypsibidae Hypsihius conoergens (Urbanowicz, 1925) Hypsihrus dujardini (Doyere, 1840) Isohypsibius 5p. nov. Diphuscon mirubile Dastych, 1984 Diphuscon pingue (Marcus, 1936) Macrobiotidae Macrohiotus furciger Murray, 1906 Macrohiotus hufelundi Schultzc, 1843 Macrobiotus lioiue Ramazzotti, 1962 Macrohiofus sp. nov. Ductylohiotus dispar (Murray, 1907)

1
-

--

- -

x 1 -

xx
-

- - - - - - _ - - _ _ - - - - - 1 - _ - I - - - - - __
-

- -

- -

xx xx
5 1

xx-

xx
-

x x - - - - - xx-xx-

- x x -

xx
-

xx xx xx
-

-_

xx
-

__

xx
-

rather than in the Antarctic. In this respect the distribution of tardigrades would be similar to that of the South Georgian vascular plants (Greene 1964; Moore 1968) and the freeliving

terrestrial arthropods (Gressitt 1970), since most species are found also on the Falkland Islands or in South America, but few in the Antarctic. According to R. M ~ b j e r g Kristensen (pers.

Tuhle 3. A survey of thc tardigrades recorded from South Georgia. X = present, - = absent.

Richters
(1920)

Jennings ( 1976b)

Dastych

This study

(1 984)

Echiniscidae Mopsechiniscus imherbis (Richters, 1920) Orecllidac Oreellu sp.? Hypsibidae Hypsibius unrurcticw Richters, 1904 Hypsibius arcticus Murray. 1910 Hypsihius conoergens (Urbanowicz, 1925) Hypsibius dujurdini (Doyi.re, 1840) Hypsihius pullidus Thulin, 1911 fsohypsibius usper (Murray, 1906) fsuhypsihius prososfomus Thulin, 1928 lsohypsihius sp. nov. Rumuzzottius oberhueuseri (Doyere, 1840) Diphascon rniruhile Dastych, 19x4 Diphascon pingue (Marcus, 1936) Diphascon scoricum (Murray, 1905) Macrobiotidae Murrohiotuy furciger Murray, 1906 Macrohiofus hufelundi Schultzc, 1843 Macrohiotus liviue Ramazzotti, 1962 Mucrohiotw sp. nov. Dactylobiotus dispur (Murray, 1907) Milnesiidae Milnesium fardigrudum Doyere, 1840

X
X2

X
-

X X X
X X X

The following names were used by Richters (1920): I: Echiniscus imherhis, 2 : Echiniscus macronyx (present status unclear, probably an Oreella sp.), Macrohiotus murruyi, : Macrohiotus asper, 5 : Macrobiotus furcutus.

294 P. S. Ottesen & T. Meier

comm.), the tardigrade fauna of South Georgia show similarities to that of Chile and Argentina. This similarity of the biota is probably a result of geological history and the prevailing winds which blow from the west. However, a major difference between the composition of terrestrial arthropods and vascular plants and tardigrades is the large amount of cosmopolitan or widespread tardigrade species. In this respect the tardigrade distribution of South Georgia is more like the macrolichen distribution. Lindsay (1974) calculated that 60% of the South Georgian macrolichen species are bipolar or cosmopolitan, compared with 17% for vascular plants (Greene 1964).
Acknowledgements. - Richard Ring also sampled tardigradcs in thc Husvik arca during the 1988expcdition. Dccdec Kathman cxtractcd the eight moss samplcs and identified the tardigradcs. Thc samplcs contained some of the species found by us, but no ncw oncs. We thank thcm for thcir gcncrous cooperation and tor comments to the manuscript. We are grateful to S. Mclnnes for valuable corrections to the manuscript and information on antarctic tardigrades, K. Mllbjcrg Kristcnsen for comments and for idcntifying and checking parts of the material, W. Block and L. Sommc for corrections on style and language, the British Antarctic Survey for logistic and financial support and the Norwcgian Research Council for Scicnce and thc Humanitics (NAVF) for a grant.
Evcritt, D. A . 1981: An ecological study of an Antarctic freshwater pole. Hydrobiologia 83, 225-237. Grecne, S . W. 1964: The vascular Rora of South Gcorgia. Br. Anturct. Suro. Sci. Rep. 45, 1-58, 31 maps, 6 pls. Grcssitt, J. L. 1970: Subantarctic entomology and biogcography. Pucif. Ins. Monogr. 2.7, 295-374. Iharos. G. 1963:The zoological results of Gy. Topals collccting in South Argentina. Ann. Hist. - Nut. Mus. Nutionali.~ Hungar. 55. 293-299. Jennings, P. G. 1976a: Thc Tardigrada of Signy Islmd. South Orkncy Islands, with a note on the Rotifera. Rr. Anturct. Suro. Bull. 44, 1-25. Jennings, P. G. 1976b: Tardigrada from the Antarctic Peninsula and Scotia Ridge region. Br. Anturct. Suro. BulL 44, 77-95. Lindsay, D. C. 1974: The macrolichens of South Georgia. Br. Anturct. Surv. Sci. Rep. 89, 1-91, 3 pls. Mclnnes, S . J . & Ellis-Evans, J . C. 1987: Tardigradcs from maritime Antarctic freshwater lakes. Pp. I1 1-123 in Bertolani, R. (ed.): Biology of Turdigrades. Selected Symposia and Monographs U.2.1. 1. Mucchi, Modena. Manicdrdi. G. C. & Bcrtolani, R. 1987: First contribution to thc knowlcdgc of alpine grassland tardigradcs. Pp. 177-1 8S in Bcrtolani, R. (cd.): Biology of Turdigrades. Selected .Yyrnposiu and Monogruphs U . Z . I . 1. Mucchi, Modcna. MihclEiE. F. 1967: Ein Beitrag zur Kenntnis der Tardigraden Argentinicns. Ver. 2001. Bot. Gesel1.sch.. Wien 107, 43-56. Miller, J . D., Hornc, P., Heatwole, H., Miller, W. R. & Bridges, L. 1988: A survey of thc terrestrial Tardigrada of the Vestfold Hills, Antarctica. Hydrobiologiu 165. 197-208. Moore, D. M. 1968: The vascular flora of the Falkkand Islands. Br. Anturct. Suro. Sci. Rep. 60. 1-202, 6 pls. Morgan, C. 1. &King, P. E. 1976 BritishTardigrades. Synopsis of the British Fauna 9. Academic Press, London. 133 pp. Murray, J. 1YO6: Scottish National Antarctic Expedition. Tardigrada of the South-Orkneys. Truns. R. Soc. Edinburgh 45. 32>334. Murray, J . 1910: Tardigrada. Brit. Anturct. Exped. 190749. London l ( 5 ) . 83-185. Ramazzotti, G. & Maucci, W. 1983: II Philum Tardigrada (3rd cd.). Mem. dlstit. ltuliuno di Idrobiologiu. 1012 pp. Richtcra, F. 1920: Moosbcwohner. Win. Ergebn. Schwed. Siidpolur-Exped. 1901-1903. Stockholm 6(2). 1-16, 1 pl. Usher, M. B. & Dastych, 1. 1987: Tardigrada from the Maritime Antarctic. Br. Antarct. Suro. Bull. 77, 163-366.

References
Block, W.. S@mmc,L., Ring, R., Ottcscn. P. & Worland, M. K. 1988: Adaptation\ of arthropods to the sub-Antarctic environment. Br. Anturct. Suru. Bull. 81, 65-67. Dastych, 11. 1984: Thc Tardigrada from Antarctica with dcscriotion of scvcral ncw specles. Actu 2001. Cracou. 27, 317-436.