(Rosaceae) at Kerguelen (South Indian Ocean)
some appeared to multiply by a budding process. The identification of several hundred pure isolates is in progress. The present work was supported by National Science Foundation grant DPP 83-14180 to E.I. Friedmann, by National Aeronautics and Space Administration grant 7337 to E. I. Friedmann, and by a grant from the Deutsche Forschungsgemeinschaft to P. Hirsch. The Department of Biological Science, Florida State University, Tallahassee, kindly provided laboratory facilities to P. Hirsch during a sabbatical leave. Skillful technical assistance of B. Hoffman and A. Stutzel-Studt is gratefully acknowledged.
References Atlas, R. M., M. di Menna, and R. E. Cameron. 1978. Ecological investigations of yeasts in Antarctic soils. In B. Parker (Ed.), Terrestrial biology Ill. (Antarctic Research Series, Vol. 30.) Washington, D.C.: American Geophysical Union. Boyd, W.L., J.T. Staley, and J.W. Boyd. 1966. Ecology of soil mircoorganisms of Antarctica. In I.C.F. Tedrow (Ed.), Antarctic soils and soil forming processes. (Antarctic Research Series, Vol. 8.) Washington, D.C.: American Geophysical Union. Cameron, R.E. 1972. Microbial and ecologic investigations in Victoria Valley, southern Victoria Land, Antarctica. In G.A. Llano (Ed.), Antarctic terrestrial biology. (Antarctic Research Series, Vol. 20.) Washington, D.C.: American Geophysical Union. Cameron, R.E. 1973. Antarctic soil microbial and ecological investigations. In L.O. Quam (Ed.), Research in the Antarctic. Washington, D.C.: American Association for the Advancement of Science.
Variability in vegetative and reproductive biomass in Acaena adscendens VahI. (Rosaceae) at Kerguelen (South Indian Ocean) J.M. LAWRENCE and J.B. MCCLINTOCK* Department of Biology University of South Florida Tampa, Florida 33620
The rosaceus herb Acaena adscendens currently covers much of the eastern part of the Courbet Peninsula on the large island of Grande Terre at Kerguelen. There is a general consensus that A. adscendens has extended its distribution on the island as a result of the elimination of its competitors following the release of rabbits there by the British Transit of Venus Expedition in 1874 (Chastain 1958; Cours 1958, 1959; Aubert de la Rile 1964). A. adscendens was the only species on the island that possessed a radicular system allowing it to recover from winter herbivory. Chastain, however, suggested some caution in attributing the emergence of A. adscendens as the dominant plant solely to the 186
Cameron, R.E., J . King, and C.N. David. 1970. Microbiology, ecology and microlimnology of soil sites in Dry Valleys of southern Victoria Land, Antarctica. In M.W. Holdgate (Ed.), Antarctic ecology (Vol. 2). London: Academic Press. Friedmann, E. 1982. Endolithic microorganisms in the Antarctic cold desert. Science, 215, 1045 - 1053. Gallikowski, C. A. 1985. The investigation of microorganisms in antarctic soil and their correlation. (Masters thesis, University of Kiel, Federal Re-
public of Germany.) (In German.) Hirsch, P., and E.I. Friedmann. 1984. Endolithic, oligotrophic microorganisms from rock samples and soils of the Dry Valleys, McMurdo, Antarctica. Abstracts, American Society of Microbiologists, 102. Horowitz, N.H., A.J. Bauman, R.E. Cameron, P.J. Geiger, J.S. Hubbard, G.P. Shulman, P.G. Simmonds, and K. Westberg. 1969. Sterile soils from Antarctica: Organic analysis. Science, 164, 1054 - 1056. Horowitz, N.H., R. Cameron, and J. S. Hubbard. 1972. Microbiology of the Dry Valleys of Antarctica. Science, 176, 242 - 245. Johnson, R.M., J.M. Madden, and J.R. Swafford. 1978. Taxonomy of Antarctic bacteria from soils and air primarily of the McMurdo Station and Victoria Land Dry Valleys region. In B. Parker (Ed.), Terrestrial biology III. (Antarctic Research Series, Vol. 30.) Washington, D.C.: American Geophysical Union. Staley, J.T. 1968. Prosthecomicrohium and Ancalomicrohium, new prosthecate fresh water bacteria. Journal of Bacteriology, 95, 1922 - 1948. Uydess, IL., and W. Vishniac. 1976. Electron microscopy of Antarctic soil bacteria. In M.R. Heinrich (Ed.), Extreme environments. Mechanisms of microbial adaptations. London: Academic Press. Vishniac, H.S. 1982. An enation system for the isolation of Antarctic yeasts inhibited by conventional media. Canadian Journal of Microbiology, 29, 90 - 95. Vishniac, H.S., and W.P. Hempfling. 1979. Evidence of an indigenous microbiota (yeasts) in the Dry Valleys of Antarctica. Journal of General Microbiology, 112, 301 - 314.
presence of rabbits, noting that the ascendency of A. adscendens over Azorella selago Hooker (Ombelliferae) began before the introduction of rabbits. He hypothesized that physical factors might be involved. There is some support for this in the observations of Schalke and van Zinderen Bakker (1971) of an alteration in the occurrence of the pollen of A. adscendens and A. se/ago in cores taken on Marion Island. They interpreted these alternating phases as an indication of alternating climatic changes. Chastain (1958) noted that A. adscendens existed in two major growth forms on Courbet Peninsula. One form was characterized by long woody stems and roots and large leaves. This form occurred in meadows where there was a relatively high level of water (average of 72 percent), an average pH of 5.5, and a high level of colloidal organic material. This form also occurred in heaths where the water level was lower (average of 27.7 percent), the average pH was higher (7.01), and the soil was a brown-black humus with sand and pebbles. Although these forms were similar in both habitats, plants in the meadows had a more luxuriant vegetative growth and did not bloom, while plants in the heath did bloom despite much less vegetative growth. The second form of A. adscendens occurred in areas with high levels of water (70 to 90 percent) and a pH of 6.4 to 6.9. This * Present address: Center for Marine Studies, University of California, Santa Cruz, California 95064.
ANTARCTIC JOURNAL
form was slender and creeping, scrawny in appearance, and with few leaves. Chastain made no quantitative measurements of the different forms. In January 1982, we collected A. adscendens from various populations in the vicinity of Port-aux-Francais. The total biomass of the individual plants and of their structural components was measured. We also counted the number of blossoms to indicate the fecundity of the plants. A. adscendens from the different populations differed distinctly in the biomass allocated to various structural components, the total biomass, and the number of blossoms (table). Plants with the highest level of vegetative growth (site 3) and those with the lowest level of vegetative growth (site 4) were not reproductive. The lack of reproduction in the former implies that total allocation of production to vegetative growth is adaptive under certain situations, and not due to inadequate material or energy. There must be internal control of allocation in such situations. The lack of reproduction in the latter is probably due to inadequate production in unfavorable conditions of high exposure and low soil water. Reproduction by A. adscendens occurs in those locations where production is sufficient for reproduction (sites 1 and 2), but not so great that reproduction is completely inhibited (site 3). Characteristic of different populations of Acaena adscendens on Grande Terre, Kerguelen, in January 1982 Site
Biomass (grams wet weight per square meter) Leaves and stems 960 608 2496 208 Blossoms and blossom stalks 272 112 0 0 1232 720 2496 208 Total Number of blossoms (per square meter) Exposure to wind Soil water (%)
1985 REVIEW
272 144 0 0 Moderate High Low High 87 38 65 18
Boulière (1982) noted that there have been few estimations of biomass and production of plants on the subantarctic islands, and specifically alluded to the lack of such data for Kerguelen and Crozet Islands. Because the leaves of A. adscendens completely die back in the winter (Cours 1958; Aubert de La Rile 1964) and growth generally stops at the end of February, the biomass reported here is an approximate estimation of annual production of A. adscendens on Kerguelen. Proximate analysis of the various components of the plant will allow us to calculate the amounts of soluble protein, soluble carbohydrate, fiber, and joules present in individual plants and the populations. This will indicate the manner in which total production is divided among these organic classes, and will have implications for understanding the way in which rabbits on Kerguelen use A. adscendens as food. This research was supported by National Science Foundation grant DPP 81-08992 and by the administration of the Terres Australes et Antarctiques Françaises. We thank C.J. Dawes for commenting on the manuscript.
References Aubert de la Rue, E. 1964. Observations on the characters and the distribution of the vegetation on the Kerguelen Islands. Comitë National Francaise des Recherches Antarctiques, (10), 1 - 60. (In French.) Boulière, F. 1982. Definition of the objectives of the meeting. Comitd National Fran çaise des Recherches Antarctiques, (51), 513 - 516. (In French.) Chastain, A. 1958. The flora and vegetation of the Kerguelen Islands: Polymorphism of southern species. Memoires du Museum National d'Histoire Naturelles. Series B, 11(1), 1 - 136. (In French.) Cours, P. 1958. Regarding the flora of the Archipelago of Kerguelen. Revue Territoire Australes et Antarctiques Françaises, (4 & 5), 10 - 32. (In French.) Cours, P. 1959. Flora and vegetation of the Archipelago of Kerguelen. Revue Territoire Australes et Antarctiques Francaises, (8 & 9), 3 - 40. (In French.) Schalke, H.J.C., and E. H. van Zinderen Bakker, Sr. 1971. History of the vegetation. In J.M. Winterbottom, and R.A. Dyer (Eds.), Marion and Prince Edward Islands. Report on the South African Biological and Geological Expedition, 1965 - 1966. Cape Town: A.A. Balkema.
187
References Atlas, R. M., M. di Menna, and R. E. Cameron. 1978. Ecological investigations of yeasts in Antarctic soils. In B. Parker (Ed.), Terrestrial biology Ill. (Antarctic Research Series, Vol. 30.) Washington, D.C.: American Geophysical Union. Boyd, W.L., J.T. Staley, and J.W. Boyd. 1966. Ecology of soil mircoorganisms of Antarctica. In I.C.F. Tedrow (Ed.), Antarctic soils and soil forming processes. (Antarctic Research Series, Vol. 8.) Washington, D.C.: American Geophysical Union. Cameron, R.E. 1972. Microbial and ecologic investigations in Victoria Valley, southern Victoria Land, Antarctica. In G.A. Llano (Ed.), Antarctic terrestrial biology. (Antarctic Research Series, Vol. 20.) Washington, D.C.: American Geophysical Union. Cameron, R.E. 1973. Antarctic soil microbial and ecological investigations. In L.O. Quam (Ed.), Research in the Antarctic. Washington, D.C.: American Association for the Advancement of Science.
Variability in vegetative and reproductive biomass in Acaena adscendens VahI. (Rosaceae) at Kerguelen (South Indian Ocean) J.M. LAWRENCE and J.B. MCCLINTOCK* Department of Biology University of South Florida Tampa, Florida 33620
The rosaceus herb Acaena adscendens currently covers much of the eastern part of the Courbet Peninsula on the large island of Grande Terre at Kerguelen. There is a general consensus that A. adscendens has extended its distribution on the island as a result of the elimination of its competitors following the release of rabbits there by the British Transit of Venus Expedition in 1874 (Chastain 1958; Cours 1958, 1959; Aubert de la Rile 1964). A. adscendens was the only species on the island that possessed a radicular system allowing it to recover from winter herbivory. Chastain, however, suggested some caution in attributing the emergence of A. adscendens as the dominant plant solely to the 186
Cameron, R.E., J . King, and C.N. David. 1970. Microbiology, ecology and microlimnology of soil sites in Dry Valleys of southern Victoria Land, Antarctica. In M.W. Holdgate (Ed.), Antarctic ecology (Vol. 2). London: Academic Press. Friedmann, E. 1982. Endolithic microorganisms in the Antarctic cold desert. Science, 215, 1045 - 1053. Gallikowski, C. A. 1985. The investigation of microorganisms in antarctic soil and their correlation. (Masters thesis, University of Kiel, Federal Re-
public of Germany.) (In German.) Hirsch, P., and E.I. Friedmann. 1984. Endolithic, oligotrophic microorganisms from rock samples and soils of the Dry Valleys, McMurdo, Antarctica. Abstracts, American Society of Microbiologists, 102. Horowitz, N.H., A.J. Bauman, R.E. Cameron, P.J. Geiger, J.S. Hubbard, G.P. Shulman, P.G. Simmonds, and K. Westberg. 1969. Sterile soils from Antarctica: Organic analysis. Science, 164, 1054 - 1056. Horowitz, N.H., R. Cameron, and J. S. Hubbard. 1972. Microbiology of the Dry Valleys of Antarctica. Science, 176, 242 - 245. Johnson, R.M., J.M. Madden, and J.R. Swafford. 1978. Taxonomy of Antarctic bacteria from soils and air primarily of the McMurdo Station and Victoria Land Dry Valleys region. In B. Parker (Ed.), Terrestrial biology III. (Antarctic Research Series, Vol. 30.) Washington, D.C.: American Geophysical Union. Staley, J.T. 1968. Prosthecomicrohium and Ancalomicrohium, new prosthecate fresh water bacteria. Journal of Bacteriology, 95, 1922 - 1948. Uydess, IL., and W. Vishniac. 1976. Electron microscopy of Antarctic soil bacteria. In M.R. Heinrich (Ed.), Extreme environments. Mechanisms of microbial adaptations. London: Academic Press. Vishniac, H.S. 1982. An enation system for the isolation of Antarctic yeasts inhibited by conventional media. Canadian Journal of Microbiology, 29, 90 - 95. Vishniac, H.S., and W.P. Hempfling. 1979. Evidence of an indigenous microbiota (yeasts) in the Dry Valleys of Antarctica. Journal of General Microbiology, 112, 301 - 314.
presence of rabbits, noting that the ascendency of A. adscendens over Azorella selago Hooker (Ombelliferae) began before the introduction of rabbits. He hypothesized that physical factors might be involved. There is some support for this in the observations of Schalke and van Zinderen Bakker (1971) of an alteration in the occurrence of the pollen of A. adscendens and A. se/ago in cores taken on Marion Island. They interpreted these alternating phases as an indication of alternating climatic changes. Chastain (1958) noted that A. adscendens existed in two major growth forms on Courbet Peninsula. One form was characterized by long woody stems and roots and large leaves. This form occurred in meadows where there was a relatively high level of water (average of 72 percent), an average pH of 5.5, and a high level of colloidal organic material. This form also occurred in heaths where the water level was lower (average of 27.7 percent), the average pH was higher (7.01), and the soil was a brown-black humus with sand and pebbles. Although these forms were similar in both habitats, plants in the meadows had a more luxuriant vegetative growth and did not bloom, while plants in the heath did bloom despite much less vegetative growth. The second form of A. adscendens occurred in areas with high levels of water (70 to 90 percent) and a pH of 6.4 to 6.9. This * Present address: Center for Marine Studies, University of California, Santa Cruz, California 95064.
ANTARCTIC JOURNAL
form was slender and creeping, scrawny in appearance, and with few leaves. Chastain made no quantitative measurements of the different forms. In January 1982, we collected A. adscendens from various populations in the vicinity of Port-aux-Francais. The total biomass of the individual plants and of their structural components was measured. We also counted the number of blossoms to indicate the fecundity of the plants. A. adscendens from the different populations differed distinctly in the biomass allocated to various structural components, the total biomass, and the number of blossoms (table). Plants with the highest level of vegetative growth (site 3) and those with the lowest level of vegetative growth (site 4) were not reproductive. The lack of reproduction in the former implies that total allocation of production to vegetative growth is adaptive under certain situations, and not due to inadequate material or energy. There must be internal control of allocation in such situations. The lack of reproduction in the latter is probably due to inadequate production in unfavorable conditions of high exposure and low soil water. Reproduction by A. adscendens occurs in those locations where production is sufficient for reproduction (sites 1 and 2), but not so great that reproduction is completely inhibited (site 3). Characteristic of different populations of Acaena adscendens on Grande Terre, Kerguelen, in January 1982 Site
Biomass (grams wet weight per square meter) Leaves and stems 960 608 2496 208 Blossoms and blossom stalks 272 112 0 0 1232 720 2496 208 Total Number of blossoms (per square meter) Exposure to wind Soil water (%)
1985 REVIEW
272 144 0 0 Moderate High Low High 87 38 65 18
Boulière (1982) noted that there have been few estimations of biomass and production of plants on the subantarctic islands, and specifically alluded to the lack of such data for Kerguelen and Crozet Islands. Because the leaves of A. adscendens completely die back in the winter (Cours 1958; Aubert de La Rile 1964) and growth generally stops at the end of February, the biomass reported here is an approximate estimation of annual production of A. adscendens on Kerguelen. Proximate analysis of the various components of the plant will allow us to calculate the amounts of soluble protein, soluble carbohydrate, fiber, and joules present in individual plants and the populations. This will indicate the manner in which total production is divided among these organic classes, and will have implications for understanding the way in which rabbits on Kerguelen use A. adscendens as food. This research was supported by National Science Foundation grant DPP 81-08992 and by the administration of the Terres Australes et Antarctiques Françaises. We thank C.J. Dawes for commenting on the manuscript.
References Aubert de la Rue, E. 1964. Observations on the characters and the distribution of the vegetation on the Kerguelen Islands. Comitë National Francaise des Recherches Antarctiques, (10), 1 - 60. (In French.) Boulière, F. 1982. Definition of the objectives of the meeting. Comitd National Fran çaise des Recherches Antarctiques, (51), 513 - 516. (In French.) Chastain, A. 1958. The flora and vegetation of the Kerguelen Islands: Polymorphism of southern species. Memoires du Museum National d'Histoire Naturelles. Series B, 11(1), 1 - 136. (In French.) Cours, P. 1958. Regarding the flora of the Archipelago of Kerguelen. Revue Territoire Australes et Antarctiques Françaises, (4 & 5), 10 - 32. (In French.) Cours, P. 1959. Flora and vegetation of the Archipelago of Kerguelen. Revue Territoire Australes et Antarctiques Francaises, (8 & 9), 3 - 40. (In French.) Schalke, H.J.C., and E. H. van Zinderen Bakker, Sr. 1971. History of the vegetation. In J.M. Winterbottom, and R.A. Dyer (Eds.), Marion and Prince Edward Islands. Report on the South African Biological and Geological Expedition, 1965 - 1966. Cape Town: A.A. Balkema.
187
