Smithsonian at the Poles: Contributions to International Polar

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invertebrates were placed in the bag and sampled at different time intervals. We have been successful at culturing larvae of the clam Laternula eliptica for over 13 months under the ice, and recovering fully metamorphosed juveniles at the end of that time period. The ability to place embryos and larvae under the ice in culture containers in the Austral summer, then leave them in place through the winter season, means that we now have the potential to work with the full lifecycle of some marine invertebrate larvae. Because of the slow developmental rates of these larvae and protracted lifespans, there are relatively few studies that have been able to collect any data on the later life-stage as they approach metamorphosis. However, divers can now establish cultures in situ under the ice, then return at any time point later on to sample individuals. This increase in the time span for which we can now study will fi ll in the large gap in our current knowledge of what happens at the end of the Austral winter period when the larvae are ready to become juveniles. In order to understand how larvae are adapted to survive in polar environments, we really need to make our best measurements and execute our most exact experiments on the individuals that have survived for the complete developmental period and are now ready to become juveniles. If only 10 percent of a cohort survives to this stage (�12 months development), then making early measurements on the other 90% (at 1 month) that were destined to die would essentially just give you information about what does not work. It is the survivors that hold the key to understanding how these organisms are adapted to persist in a harsh polar environment. In essence, all the existing studies on adaptations in polar marine larvae that have made measurements from bulk cultures at early developmental time points could be grossly misleading. All of those individuals are not likely to survive the full year to recruitment. Consequently, we need an experimental culturing system that will allow scientists to work with larvae that have survived the harsh polar environment. Those are the individuals that have the key to understanding adaptive processes. Overall, the in situ culturing approach offers us three main advantages: 1. Large numbers of individuals can be cultured with relatively little husbandry effort. Once the culture containers are setup and stocked, then the only effort necessary is for a dive team to periodically sample and remove individuals. There is no feeding and little maintenance required. COLD ADAPTATION IN POLAR MARINE INVERTEBRATES 261 2. Larvae can be cultured under very natural conditions without laboratory artifacts. The most important variable to control is temperature and by not having open culture containers in an aquarium room, there is no worry about the temperature in the vessels changing because of problems with electricity supply, pumps breaking down and losing the cold sea water supply rate, or someone just changing the thermostat within the aquarium room. The second most important variable is food, and under in situ conditions, the feeding larvae will receive a diet of natural species and in a natural supply. 3. Long term cultures can be maintained across the entire developmental period, which in most polar marine invertebrates can easily extend upwards of a year. This approach will provide access to larvae that have successfully survived their full lifecycle in a harsh polar environment. CONCLUSION The S. neumayeri distributions in developmental rate (Figure 3), respiration distributions (Figure 4), transcriptome profi les (Figure 5), and gene methylation (data not shown) have focused our attention on trying to understand the functional signifi cance of interindividual variability at these levels of biological organization. In a lifehistory model that selects for a prolonged larval lifespan, it is intriguing to ask whether or not it is a reduction in individual metabolic rates (Figure 2A) or an increase in the cohort variance in metabolic rates (Figure 2B) that could account for the adaptation in metabolic phenotypes. The metabolic lifespans of polar invertebrate larvae could be under the same genetic determinants as other temperate species, but changes in patterns of gene regulation could substantially alter the distribution of physiological phenotypes within a cohort. Being able to study long-lived larvae that are ready to become juveniles holds the key for deciphering the adaptive mechanism that may be operative at the level of a full cohort to ensure that some percentage is capable of surviving. Selection is surely not operating to force the survival function of all individuals within a cohort. Only enough need to survive to keep a population established and stable. Scientifi c diving will be an important component of discovering how these animals are adapted to survive. The opportunity to now work in situ with embryos and larvae will open new avenues of research and understanding. Even though the under ice work is not complex, it is nonetheless
262 SMITHSONIAN AT THE POLES / MARSH rigorous and demanding. Any diver working on the bottom for more than 45 minutes will readily attest to the extreme nature of the cold that impacts all organisms in that environment. Being in the water gives one a unique perspective on the survival challenges that are facing the embryos and larvae of these polar marine invertebrates. ACKNOWLEDGMENTS This work was conducted with the help, collaboration, and participation of numerous students and postdoctoral associates in my lab and I am grateful for their efforts, energies and cold tolerance. The research was supported by a grant from the National Science Foundation, Offi ce of Polar Programs (#02– 38281 to AGM). LITERATURE CITED Buckley, B. A., S. P. Place, and G. E. Hofmann. 2004. Regulation of Heat Shock Genes in Isolated Hepatocytes from an Antarctic Fish, Trematomus bernacchii. Journal of Experimental Biology, 207: 3649– 3656. Cheng, C. H. C., P. A. Cziko, and E. W. Clive. 2006. Nonhepatic Origin of Notothenioid Antifreeze Reveals Pancreatic Synthesis as Common Mechanism in Polar Fish Freezing Avoidance. Proceedings of the National Academies of Science, 103: 10491– 10496. Clarke, A. 1991. What Is Cold Adaptation and How Should We Measure It? American Zoologist, 31: 81– 92. Clarke, A., N. M. Johnston, E. J. Murphy, and A. D. Rogers. 2007. Introduction: Antarctic Ecology from Genes to Ecosystems: The Impact of Climate Change and the Importance of Scale. Philosophical Transactions of the Royal Society B-Biological Sciences, 362: 5– 9. Detrich, H. W., S. K. Parker, R. C. Williams, E. Nogales, and K. H. Downing. 2000. Cold Adaptation of Microtubule Assembly and Dynamics— Structural Interpretation of Primary Sequence Changes Present in the Alpha- and Beta-Tubulins of Antarctic Fishes. Journal of Biological Chemistry, 275: 37038– 37047. Devries, A. L. and C. H. C. Cheng. 2000. Freezing Avoidance Strategies Differ in Antarctic And Arctic Fishes. American Zoologist, 40: 997. Eastman, J. T. 2005. The Nature of the Diversity of Antarctic Fishes. Polar Biology, 28: 93– 107. Fraser, K. P. P., L. S. Peck, and A. Clarke. 2004. Protein Synthesis, RNA Concentrations, Nitrogen Excretion, and Metabolism Vary Seasonally in the Antarctic Holothurian Heterocucumis steineni (Ludwig 1898). Physiological and Biochemical Zoology, 77: 556– 569. Fraser, K. P. P., and A. D. Rogers. 2007. Protein Metabolism in Marine Animals: The Underlying Mechanism of Growth. Advances in Marine Biology, 52: 267– 362. Giaever, G., A. M. Chu, L. Ni, C. Connelly, L. Riles, S. Veronneau, S. Dow, A. Lucau-Danila, K. Anderson, B. Andre, A. P. Arkin, A. Astromoff, M. El Bakkoury, R. Bangham, R. Benito, S. Brachat, S. Campanaro, M. Curtiss, K. Davis, A. Deutschbauer, K. D. Entian, P. Flaherty, F. Foury, D. J. Garfi nkel, M. Gerstein, D. Gotte, U. Guldener, J. H. Hegemann, S. Hempel, Z. Herman, D. F. Jaramillo, D. E. Kelly, S. L. Kelly, P. Kotter, D. LaBonte, D. C. Lamb, N. Lan, H. Liang, H. Liao, L. Liu, C. Y. Luo, M. Lussier, R. Mao, P. Menard, S. L. Ooi, J. L. Revuelta, C. J. Roberts, M. Rose, P. Ross-Macdonald, B. Scherens, G. Schimmack, B. Shafer, D. D. Shoemaker, S. Sookhai-Mahadeo, R. K. Storms, J. N. Strathern, G. Valle, M. Voet, G. Volckaert, C. Y. Wang, T. R. Ward, J. Wilhelmy, E. A. Winzeler, Y. H. Yang, G. Yen, E. Youngman, K. X. Yu, H. Bussey, J. D. Boeke, M. Snyder, P. Philippsen, R. W. Davis, and M. Johnston. 2002. Functional Profi ling of the Saccharomyces cerevisiae Genome. Nature, 418: 387– 391. Haaf, T. 2006. Methylation Dynamics in the Early Mammalian Embryo: Implications of Genome Reprogramming Defects for Development. Current Topics in Microbiology and Immunology, 310: 13– 22. Hofmann, G. E., S. G. Lund, S. P. Place, and A. C. Whitmer. 2005. Some Like It Hot, Some Like It Cold: The Heat Shock Response Is Found in New Zealand but Not Antarctic Notothenioid Fishes. Journal of Experimental Marine Biology and Ecology, 316: 79– 89. Hoover, C. A., M. Slattery, and A. G. Marsh. 2007a. A Functional Approach to Transcriptome Profi ling: Linking Gene Expression Patterns to Metabolites That Matter. Marine Biotechnology, doi:10.1007/s10126– 007– 9008– 2. ———. 2007b. Comparing Gene Expression Profi les of Benthic Soft Coral Species, Sinularia polydactyla, S. maxima, and Their Putative Hybrid. Comparative Biochemistry and Physiology (Part D), 2: 135– 143. ———. 2007c. Profi ling Transcriptome Complexity during Secondary Metabolite Synthesis Induced by Wound Stress in a Benthic Soft Coral, Sinularia polydactyla. Marine Biotechnology, doi:10.1007/ s10126– 006– 6048-y. Huening, M. A., D. W. Sevilla, A. Potti, D. H. Harpole, J. R. Nevins, and M. B. Datto. 2006. A First Step towards Validating Microarray Gene Expression Profi ling for Clinical Testing: Characterizing Biological Variance. Journal of Molecular Diagnostics, 8: 660– 665. Kaminsk, Z., S. C. Wang, S. Ziegler, I. Gottesman, A. Wong, A. McRae, P. Visscher, N. Martin, and A. Petronis. 2006. Epigenomic Microarray Profi ling of DNA Methylation Variability in Monozygotic and Dizygotic Twins. American Journal of Medical Genetics Part B-Neuropsychiatric Genetics, 141B: 708– 714. Kendall, L. R., E. McMullin, and A. G. Marsh. 2005. Genome Methylation Patterns during Early Development in the Antarctic Sea Urchin, Sterechinus neumayeri. Integrative and Comparative Biology, 45: 1153. King, M. C., and A. C. Wilson. 1975. Evolution at 2 Levels in Humans and Chimpanzees. Science, 188: 107– 116. Levenson, J. M., and J. D. Sweatt. 2006. Epigenetic Mechanisms: A Common Theme in Vertebrate and Invertebrate Memory Formation. Cellular and Molecular Life Sciences, 63: 1009– 1016. MacKay, A. B., A. A. Mhanni, R. A. McGowan, and P. H. Krone. 2007. Immunological Detection of Changes in Genomic DNA Methylation during Early Zebrafi sh Development. Genome, 50: 778– 785. Marsh, A. G., S. Cohen, and C. E. Epifanio. 2001a. Larval Energy Metabolism and Physiological Variability in the Shore Crab, Hemigrapsus sanguineus. Marine Ecology Progress Series, 218: 303– 309. Marsh, A. G., and K. T. Fielman. 2005. Transcriptome Profi ling of Individual Larvae of Two Different Developmental Modes in the Poecilogonous Polychaete Streblospio benedicti (Spionidae). Journal of Experimental Zoology—Molecular Development and Evolution, 304B: 238– 249. Marsh, A. G., and D. T. Manahan. 1999. A Method for Accurate Measurements of the Respiration Rates of Marine Invertebrate Embryos and Larvae. Marine Ecology Progress Series, 184: 1– 10. ——— . 2000. Metabolic Differences between “Demersal” and “Pelagic” Development of the Antarctic Sea Urchin Sterechinus neumayeri. Marine Biology, 137: 215– 221. Marsh, A. G., R. E. Maxson, and D. T. Manahan. 2001b. High Macromolecular Synthesis with Low Metabolic Cost in Antarctic Sea Urchin Embryos. Science, 291: 1950– 1952. McClintock, J. B., C. D. Amsler, B. J. Baker, and R. W. M. van Soest. 2005. Ecology of Antarctic Marine Sponges: An Overview. Integrative and Comparative Biology, 45: 359– 368.
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Smithsonian at the Poles Contributi
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Contents FOREWORD by Ira Rubinoff i
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Elaina Jorgensen, Alaska Fisheries
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CONTENTS vii Watching Star Birth fr
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x SMITHSONIAN AT THE POLES blooms i
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xii SMITHSONIAN AT THE POLES Change
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xiv SMITHSONIAN AT THE POLES Museum
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Advancing Polar Research and Commun
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James P. Espy (1785- 1860), the fi
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ology fueled hopes that the scienti
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Meteorologists also provided critic
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visualize weather patterns remotely
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in ice sheets. The latest collapse
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Cooperation at the Poles? Placing t
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British Association for the Advance
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cations, in which the Smithsonian s
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ence” (Robinson, 2006: 76), he ha
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Taylor, C. J. 1981. First Internati
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24 SMITHSONIAN AT THE POLES / KORSM
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36 SMITHSONIAN AT THE POLES / De VO
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From Ballooning in the Arctic to 10
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ern Svalbard in 1896 (Capelotti, 19
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FIGURE 2. The Andrée campsite in 1
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National Zoo in Washington, D.C.—
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FROM BALLOONING TO 10,000-FOOT RUNW
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e rapidly deployed to South America
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“Of No Ordinary Importance”: Re
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1942). Ethnological collecting had
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group. More importantly, Murdoch’
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gan to study the question of Indian
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small museums and culture centers i
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fer of Alaskan objects and informat
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fi rst time in polar research— di
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Boas, Franz. 1888a. “The Central
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Lindsay, Debra. 1993. Science in th
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80 SMITHSONIAN AT THE POLES / FIENU
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90 SMITHSONIAN AT THE POLES / BURCH
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100 SMITHSONIAN AT THE POLES / CROW
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From Tent to Trading Post and Back
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in befriending an extraordinary Inu
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The Smithsonian Institution’s pre
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of departure for research during th
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FIGURE 8. A drawing of the so-calle
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situated experiential education and
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the Past: Archaeologists, Native Am
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130 SMITHSONIAN AT THE POLES / KRUP
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144 SMITHSONIAN AT THE POLES / PARK
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Brooding and Species Diversity in t
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iefl y consider below some of the i
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ment. Consequently, the selective e
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ated from the Antarctic. Strong cur
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the four main genera of brooding sc
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ing such cryptic speciation suggest
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LITERATURE CITED Absher, T. M., G.
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Madon-Senez, C. 1998. Disparité Mo
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Persistent Elevated Abundance of Oc
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ABUNDANCE OF ANTARCTIC OCTOPODS 199
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206 SMITHSONIAN AT THE POLES / WINS
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208 SMITHSONIAN AT THE POLES / WINS
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Page 240 and 241: Considerations of Anatomy, Morpholo
Page 242 and 243: Many theories have been hypothesize
Page 244 and 245: FIGURE 4. A three dimensional recon
Page 246 and 247: are wider and more bulbous in the a
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Page 252 and 253: DISCUSSION Imaging and dissection o
Page 254 and 255: out its length. The pulp chamber al
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Page 258 and 259: Scientifi c Diving Under Ice: A 40-
Page 260 and 261: TABLE 2. Principal Investigators an
Page 262 and 263: attaching ice anchors to the chunks
Page 264 and 265: dumping of weight under water. The
Page 266 and 267: MARINE LIFE HAZARDS Few polar anima
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Page 270 and 271: Environmental and Molecular Mechani
Page 272 and 273: face (�1.8°C) are likely to pose
Page 274 and 275: FIGURE 2. Illustrated selection shi
Page 276 and 277: FIGURE 4. Distribution of respirati
Page 280: Meehan, R. R., D. S. Dunican, A. Ru
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312 SMITHSONIAN AT THE POLES / SMIT
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320 SMITHSONIAN AT THE POLES / KIEB
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Capital Expenditure and Income (For
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tal breeding systems (Boyd, 1998; T
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FIGURE 3. Pup mass gain in relation
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MONITORING FOOD CONSUMPTION DURING
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primary productivity in McMurdo Sou
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Non-steady-State Systems. Journal o
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Latitudinal Patterns of Biological
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and perhaps others, may have invade
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colonizing the Arctic Ocean under c
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due in part to the great distances
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and (2) human-mediated responses to
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Lewis, P. N., M. Riddle, and C. L.
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Cosmology from Antarctica Robert W.
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There they found better observing c
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TELESCOPES AND INSTRUMENTS AT THE S
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Gaier, T., J. Schuster, and P. Lubi
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J. M. Kovac, C. L. Kuo, A. E. Lange
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370 SMITHSONIAN AT THE POLES / MART
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374 SMITHSONIAN AT THE POLES / WALK
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Watching Star Birth from the Antarc
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STAR BIRTH FROM THE ANTARCTIC PLATE
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is constructing and deploying the P
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Antarctic Meteorites: Exploring the
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at the Field Museum, and pieces wer
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the description and curation of Ant
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led these committees throughout the
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Index AAUS. See American Academy of
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temperature, 350-355 tourism, 354 B
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Fishing. See also Biological invasi
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McMurdo Station, xiv, 265-267, 393
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Rogick, Mary, 206 Ronne, Finn, 55 R
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U.S. Naval Support Force Antarctica
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