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![Copyright # 2008 by John Wiley & Sons, Inc. All rights reserved.
Published by John Wiley & Sons, Inc., Hoboken, New Jersey
Published simultaneously in Canada
Wiley-Blackwell is an imprint of John Wiley & Sons, formed by the merger of Wiley’s global Scientific,
Technical, and Medical business with Blackwell Publishing.
No part of this publication may be reproduced, stored in a retrieval system, or transmitted in any form or by
any means, electronic, mechanical, photocopying, recording, scanning, or otherwise, except as permitted
under Sections 107 or 108 of the 1976 United States Copyright Act, without either the prior written per-
mission of the Publisher, or authorization through payment of the appropriate per-copy fee to the Copyright
Clearance Center, Inc., 222 Rosewood Drive, Danvers, MA 01923, 978-750-8400, fax 978-750-4470, or
on the web at www.copyright.com. Requests to the Publisher for permission should be addressed to the
Permissions Department, John Wiley & Sons, Inc., 111 River Street, Hoboken, NJ 07030, 201-748-
6011, fax 201-748-6008, or online at http://www.wiley.com/go/permission.
Limit of Liability/Disclaimer of Warranty: While the publisher and author have used their best efforts in
preparing this book, they make no representations or warranties with respect to the accuracy or complete-
ness of the contents of this book and specifically disclaim any implied warranties of merchantability or
fitness for a particular purpose. No warranty may be created or extended by sales representatives or
written sales materials. The advice and strategies contained herein may not be suitable for your situation.
You should consult with a professional where appropriate. Neither the publisher nor author shall be liable
for any loss of profit or any other commercial damages, including but not limited to special, incidental,
consequential, or other damages.
For general information on our other products and services or for technical support, please contact
our Customer Care Department within the United States at 800-762-2974, outside the United States at
317-572-3993 or fax 317-572-4002.
Wiley also publishes its books in variety of electronic formats. Some content that appears in print may
not be available in electronic format. For more information about Wiley products, visit our web site at
www.wiley.com.
Library of Congress Cataloging-in-Publication Data:
Kirchman, David L.
Microbial ecology of the oceans / [edited by] David L. Kirchman. -- 2nd ed.
p. cm.
Includes bibliographical references and index.
ISBN 978-0-470-04344-8 (pbk.)
1. Marine microbiology. 2. Marine ecology. 3. Carbon cycle
(Biogeochemistry) I. Title.
QR106.M53 2008
5790
.177--dc22 2007051389
Printed in the United States of America
10 9 8 7 6 5 4 3 2 1](https://image.slidesharecdn.com/35822-250722083229-7d8f255a/75/Microbial-Ecology-of-the-Oceans-Second-Edition-David-L-Kirchman-8-2048.jpg)
![CONTRIBUTORS
ODED BÉJÀ Faculty of Biology, Technion–Israel Institute of Technology,
Haifa, Israel [beja@techunix.technion.ac.il]
MYA BREITBART College of Marine Science, University of South Florida,
St. Petersburg, FL 33701, U.S.A. [mya@marine.usf.edu]
MATTHEW J. CHURCH Department of Oceanography, University of Hawaii,
Honolulu, HI 96822, U.S.A. [mjchurch@hawaii.edu]
TAGE DALSGAARD National Environmental Research Institute, University of Aarhus,
DK-8600 Silkeborg, Denmark [tda@dmu.dk]
JED A. FUHRMAN Department of Biological Sciences, University of Southern
California, Los Angeles, CA 90089, U.S.A. [fuhrman@usc.edu]
JOSEP M. GASOL Institut de Ciències del Mar, CMIMA (CSIC), Passeig Marı́tim de la
Barceloneta 37–49, 08003 Barcelona, Catalunya, Spain [pepgasol@icm.csic.es]
PAUL A. DEL GIORGIO Département des Sciences Biologiques, Université du Québec
à Montréal, CP 8888, succ. Centre Ville, Montréal, Québec, Canada H3C 3P8
[del_giorgio.paul@uqam.ca]
ÅKE HAGSTRÖM Kalmar University, Sweden [ake.hagstrom@hik.se]
KLAUS JÜRGENS Leibniz Institute for Baltic Sea Research, 18119 Rostock, Germany
[klaus.juergens@io-warnemuende.de]
DAVID L. KIRCHMAN College of Marine and Earth Studies, University of Delaware,
Lewes, DE 19958, U.S.A. [kirchman@udel.edu]
RAMON MASSANA Institut de Ciències del Mar, CMIMA (CSIC), Passeig
Marı́tim de la Barceloneta 37–49, 08003 Barcelona, Catalunya, Spain
[ramonm@icm.csic.es]
MATHIAS MIDDELBOE Marine Biological Laboratory, University of Copenhagen,
DK-3000 Helsingør, Denmark [mmiddelboe@bio.ku.dk]
xvii](https://image.slidesharecdn.com/35822-250722083229-7d8f255a/75/Microbial-Ecology-of-the-Oceans-Second-Edition-David-L-Kirchman-21-2048.jpg)
![MARY ANN MORAN Department of Marine Sciences, University of Georgia, Athens,
GA 30602-3636, U.S.A. [mmoran@uga.edu]
TOSHI NAGATA Ocean Research Institute, The University of Tokyo, Tokyo, Japan
[nagata@ori.u-tokyo.ac.jp]
FABRICE NOT Evolution du Plancton et PaleOceans Laboratory, CNRS, Université
Paris 06, UMR7144, Station Biologique de Roscoff, 29682, Roscoff, Cedex
BP 74, France [not@sb-roscoff.fr]
HANS W. PAERL Institute of Marine Sciences, University of North Carolina at
Chapel Hill, Morehead City, NC 28557, U.S.A. [hans_paerl@unc.edu]
CAROL ROBINSON School of Environmental Sciences, University of East Anglia,
Norwich NR4 7TJ, U.K. [carol.robinson@uea.ac.uk]
FOREST ROHWER Department of Biology, San Diego State University, San Diego,
CA 92182, U.S.A. [frohwer@gmail.com]
EVELYN SHERR College of Oceanic and Atmospheric Sciences, Oregon State
University, Corvallis, OR 97331-5503, U.S.A. [sherre@coas.oregonstate.edu]
BARRY SHERR College of Oceanic and Atmospheric Sciences, Oregon State
University, Corvallis, OR 97331-5503, U.S.A. [sherrb@coas.oregonstate.edu]
MARCELINO T. SUZUKI Chesapeake Biological Laboratory, University of
Maryland, Center for Environmental Science, Solomons, MD 20688, U.S.A.
[suzuki@cbl.umces.edu]
BO THAMDRUP Institute of Biology, University of Southern Denmark, DK-5230
Odense M, Denmark [bot@biology.sdu.dk]
ALEXANDRA Z. WORDEN Monterey Bay Aquarium Research Institute, Moss
Landing, CA 95039, U.S.A. [azworden@mbari.org]
JONATHAN P. ZEHR Ocean Sciences Department, University of California, Santa
Cruz, CA 95064, U.S.A. [zehrj@pmc.ucsc.edu]
xviii CONTRIBUTORS](https://image.slidesharecdn.com/35822-250722083229-7d8f255a/75/Microbial-Ecology-of-the-Oceans-Second-Edition-David-L-Kirchman-22-2048.jpg)
![of the fringe of the glacier which almost surrounds the Vatna Jökull.
The existence of scratched rocks in moraines in Iceland below the
limit of the glaciers does not of necessity prove that such glaciers
have bodily advanced, as during extensive eruptions of glacial
mountains huge masses of ice frequently slip forward to
considerable distances, scratching the harder and furrowing the
softer rocks in their progress, which, upon their melting, leave large
piles of glacial débris, in no way distinguishable from a moraine
stranded upon the lower elevations.
It was blowing hard from the east with heavy rain, but upon the
west side of the mountain before us (Örœfa) the sun was shining in
the most tantalising manner, so that as we urged our horses along
the heavy sands we were fain to fancy ourselves exploring those
dazzling glaciers and snowy slopes which seemed to fascinate the
sunshine and detain it from reaching us.
We were soon under the lee of the mountains before us. Sheltered
from the wind and the storm, we could stop to admire the grand
sweep from the Örœfa to the commencement of the Skeiðará Jökull.
Looking back at Núpstað, we saw it enwrapped in gloom, the clouds
clustering round the Lómagnúpar,[1] a mountain which seems to
attract all the bad weather to Núpstað and the storm sat heavily
upon the western portion of the plain of Skeiðarár Sandr, which was
exposed to the fury of the east winds.
Crossing the river Skeiðará, we reached the Saga-famed Svínafell.
Here we stayed to refresh ourselves with the national panacea for
the ills of Icelandic travel, namely, a cup of coffee of the real
Icelandic brew! The art of making good coffee is one of the greatest
accomplishments of the fair sex here, and it is a pity it is not more
generally attained by the lady population of other countries. The
occurrence of drinkable coffee in Iceland, a good cup of it being
always to be obtained at the poorest farm, is the more remarkable,
as the coffee sold by the merchants at the various stores is never of
the best quality; but is principally the Java coffee. The grand secret
of success in this special domestic art is doubtless owing to the fact](https://image.slidesharecdn.com/35822-250722083229-7d8f255a/75/Microbial-Ecology-of-the-Oceans-Second-Edition-David-L-Kirchman-29-2048.jpg)
![our friend the priest is compensated for his dreary surroundings by
having one of the prettiest Icelandic women I have seen for his wife.
She seemed quite piqued because I could not own to thinking
Sandfell a very pretty place. Going hence, we crossed the stream of
lava and agglomerate, which I was informed resulted from the
eruption of Örœfa in 1862. This stream is a remarkable one,
inasmuch as the agglomerate has flowed down in a semi-molten
state, cotemporaneously with the lava, both being mixed together;
the agglomerate appears to preponderate, but this may be the result
of the lava being of higher specific gravity, which causes it to sink to
the bottom of the stream.
We stopped for the night at Myrum,[2] on the south-west of the
Breiðamerkr Sandr. The bóndi, like all the people of this district, was
hastening to get to Papós with his wool. We supped and breakfasted
off some birds which our host called Svartfugl. They were the nicest
birds I had ever tasted in Iceland, the meat being tender and plenty
of it, and I thought so well of this dish that I took one of the birds
away with us for our lunch on the road.
Here we hired a fresh horse, leaving Paul’s, which had contracted
a sore back, and started over the Breiðamerkr Sandr. The sands, like
the Skeiðarár Sandr are the result of the great efforts of the Örœfa
and Vatna Jökulls, more especially the part of the Vatna known as
the Breiðamerkr Jökull, which was the one whose movements we
had to examine.
The road over these sands is long and dreary, especially in such
weather as had just overtaken us. We passed an extensive
encampment of farmers, who were on their way to Papós; but,
despairing of crossing the rivers which traverse the Breiðamerkr
Sandr upon such a day with heavily laden horses, they had decided
on remaining encamped upon the little patch of grass they had
reached. About one third of the way over the Sandr we arrived at
the farm of Kvísker, which is situated upon a little oasis of grass-
land. We found it a very acceptable halting-place, and although we
were wet, we were glad to sit down and take coffee and schnapps,](https://image.slidesharecdn.com/35822-250722083229-7d8f255a/75/Microbial-Ecology-of-the-Oceans-Second-Edition-David-L-Kirchman-31-2048.jpg)
Microbial Ecology of the Oceans Second Edition David L. Kirchman
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- 5. Microbial Ecology ofthe Oceans Second Edition David L. Kirchman Digital Instant Download Author(s): David L. Kirchman ISBN(s): 9780470281840, 0470281847 File Details: PDF, 22.37 MB Year: 2008 Language: english
- 6. Microbial Ecology of theOceans
- 7. Microbial Ecology of theOceans Second Edition DAVID L. KIRCHMAN College of Marine and Earth Studies, University of Delaware
- 8. Copyright # 2008by John Wiley & Sons, Inc. All rights reserved. Published by John Wiley & Sons, Inc., Hoboken, New Jersey Published simultaneously in Canada Wiley-Blackwell is an imprint of John Wiley & Sons, formed by the merger of Wiley’s global Scientific, Technical, and Medical business with Blackwell Publishing. No part of this publication may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, recording, scanning, or otherwise, except as permitted under Sections 107 or 108 of the 1976 United States Copyright Act, without either the prior written per- mission of the Publisher, or authorization through payment of the appropriate per-copy fee to the Copyright Clearance Center, Inc., 222 Rosewood Drive, Danvers, MA 01923, 978-750-8400, fax 978-750-4470, or on the web at www.copyright.com. Requests to the Publisher for permission should be addressed to the Permissions Department, John Wiley & Sons, Inc., 111 River Street, Hoboken, NJ 07030, 201-748- 6011, fax 201-748-6008, or online at http://www.wiley.com/go/permission. Limit of Liability/Disclaimer of Warranty: While the publisher and author have used their best efforts in preparing this book, they make no representations or warranties with respect to the accuracy or complete- ness of the contents of this book and specifically disclaim any implied warranties of merchantability or fitness for a particular purpose. No warranty may be created or extended by sales representatives or written sales materials. The advice and strategies contained herein may not be suitable for your situation. You should consult with a professional where appropriate. Neither the publisher nor author shall be liable for any loss of profit or any other commercial damages, including but not limited to special, incidental, consequential, or other damages. For general information on our other products and services or for technical support, please contact our Customer Care Department within the United States at 800-762-2974, outside the United States at 317-572-3993 or fax 317-572-4002. Wiley also publishes its books in variety of electronic formats. Some content that appears in print may not be available in electronic format. For more information about Wiley products, visit our web site at www.wiley.com. Library of Congress Cataloging-in-Publication Data: Kirchman, David L. Microbial ecology of the oceans / [edited by] David L. Kirchman. -- 2nd ed. p. cm. Includes bibliographical references and index. ISBN 978-0-470-04344-8 (pbk.) 1. Marine microbiology. 2. Marine ecology. 3. Carbon cycle (Biogeochemistry) I. Title. QR106.M53 2008 5790 .177--dc22 2007051389 Printed in the United States of America 10 9 8 7 6 5 4 3 2 1
- 9. CONTENTS PREFACE xv CONTRIBUTORS xvii 1INTRODUCTION AND OVERVIEW 1 David L. Kirchman Eukaryotic Phytoplankton and Cyanobacteria 3 Photoheterotrophic Bacteria 5 Dissolved Organic Material 7 Heterotrophic Bacteria 10 Marine Archaea 13 Heterotrophic Protists 14 Nanoflagellates (2–20 mm) 14 Microzooplanktonic Protists (20–200 mm) 16 Dinoflagellates 16 Marine Fungi 16 Marine Viruses 17 N2 Fixers 18 Nitrifiers and Other Chemolithotrophs 19 Denitrifiers 20 Concluding Remarks 21 Summary 22 Acknowledgments 22 References 23 v
- 10. 2 UNDERSTANDING ROLESOF MICROBES IN MARINE PELAGIC FOOD WEBS: A BRIEF HISTORY 27 Evelyn Sherr and Barry Sherr Introduction 27 Pre-1950s: The Early Years 28 1950–1974 29 1970s–1980s 32 Improvement in Methods 32 Bacterial Abundance 32 Bacterial Activity 33 Marine Heterotrophic Protists 34 The “Microbial Loop” 36 1990–Present: The Molecular Revolution 39 Summary 40 References 41 3 BACTERIAL AND ARCHAEAL COMMUNITY STRUCTURE AND ITS PATTERNS 45 Jed A. Fuhrman and Åke Hagström Introduction 45 Major Groups of Prokaryotes in Seawater 47 “Classically” Culturable Bacteria 49 The Roseobacter Clade of Marine Alphaproteobacteria 50 Gammaproteobacteria 51 Bacteroidetes 52 Cyanobacteria 52 “Sea Water” Culturable Bacteria 55 SAR11 Cluster 55 Not-Yet-Cultured Bacteria 57 Marine Gammaproteobacterial Clusters 57 Actinobacteria 58 SAR116 Cluster 59 SAR202 59 Marine Group A 59 Marine Group B 59 Betaproteobacteria 59 Marine Archaea 60 Bacterioplankton Diversity 63 Species Concept 63 vi CONTENTS
- 11. Microdiversity 64 Components ofDiversity: Richness and Evenness 65 Community Structure: Description and Factors 67 Bottom-Up Control 68 Sideways Control 69 Top-Down Control 70 “Kill the Winner” Hypothesis 71 Temporal Variation (Days to Seasonal) 72 Short-Term Variation 72 Seasonal Variation 72 Spatial Variation 74 Microscale Patterns 74 Global Distribution 75 Latitudinal Gradient and Degree of Endemism 76 Patchiness and Large Eddies 77 Summary 79 References 80 4 GENOMICS AND METAGENOMICS OF MARINE PROKARYOTES 91 Mary Ann Moran Introduction 91 The Basics of Prokaryotic Genomics 92 Genome Sequence and Assembly 92 Finding Genes 95 Finding Operons 96 Functional Annotation 96 Tame or Wild? Pure-Culture Genomics Versus Metagenomics 100 Genomics in Marine Microbial Ecology 103 The Ecology of Genome Composition 103 Reverse Biogeochemistry: Discovery of New Ecological Processes 104 Environmental Reductionism: New Details About Recognized Processes 106 Comparative Genomics and Metagenomics 107 Future Directions 122 Summary 125 Acknowledgments 125 References 125 CONTENTS vii
- 12. 5 PHOTOHETEROTROPHIC MARINEPROKARYOTES 131 Oded Béjà and Marcelino T. Suzuki Introduction 131 Facultative Photoheterotrophy by Unicellular Cyanobacteria 132 Cyanobacteria as Facultative Heterotrophs 132 Uptake of Urea and DMSP 133 Uptake of Nucleosides and Amino Acids 134 Field Studies Using Light and Dark Incubations 135 Implications of Facultative Photoheterotrophy by Cyanobacteria 138 Marine AAnP Bacteria: Habitats and Diversity 139 Rediscovery of the Marine AAnP Bacteria 139 Diversity of AAnP Bacteria 139 Physiology of AAnP Bacteria 140 AAnP Bacterial Abundance and Ecological Significance 142 Proteorhodopsin-Containing Prokaryotes 143 Proteorhodopsin Genotypes and Taxonomic Distributions 144 Proteorhodopsin Spectral Tuning 145 Proteorhodopsin-Containing Prokaryotes: Abundance and Activity 146 Proteorhodopsin-Containing Prokaryotes: Ecological Significance 150 Summary 151 References 151 6 ECOLOGY AND DIVERSITY OF PICOEUKARYOTES 159 Alexandra Z. Worden and Fabrice Not Introduction 159 Functional Roles, Classification, and Biological Traits 162 Photoautotrophs 163 Heterotrophs and Alternative Lifestyles 170 Environmental Diversity and Molecular Phylogenetics 172 Diversity of Uncultured Populations 174 Methodological Issues for envPCR Studies 178 Distribution, Abundance, and Activities 179 Methods for Quantifying Mixed Assemblages 180 Distribution, Abundance, and Activity of Mixed Picophytoplankton Assemblages 182 Quantifying Specific Picoeukaryote Populations 186 Methodological Challenges to Quantifying Specific Populations and Resolving Dynamics 190 viii CONTENTS
- 13. Mortality, Contributions toMicrobial Food Webs, and Microbial Interactions 191 Genomic Approaches to Picoeukaryote Ecology 193 Integration of Picoeukaryotes to the Microbial Food Web: Research Directions 194 Summary 195 Acknowledgments 196 References 196 7 ORGANIC MATTER–BACTERIA INTERACTIONS IN SEAWATER 207 Toshi Nagata Introduction 207 Organic Matter Inventory and Fluxes 208 DOM–Bacteria Interactions 211 Labile Low-Molecular Weight (LMW) DOM 211 Extracellular Hydrolytic Enzymes 215 Polymeric DOM—Protein as a Model 217 Refractory DOM 220 POM–Bacteria Interactions 223 POM Continuum 223 POM Fluxes 223 POM–Mineral Interactions 229 Bacterial Community Structure and Utilization of Organic Matter 230 Future Challenges 231 Summary 232 References 232 8 PHYSIOLOGICAL STRUCTURE AND SINGLE-CELL ACTIVITY IN MARINE BACTERIOPLANKTON 243 Paul A. del Giorgio and Josep M. Gasol Introduction 243 Distribution of Physiological States in Bacterioplankton Assemblages 245 The Concept of “Physiological Structure” of Bacterioplankton Assemblages 245 Starvation, Dormancy, and Viability in Marine Bacterioplankton 246 CONTENTS ix
- 14. Describing the PhysiologicalStructure of Bacterioplankton 250 Single-Cell Properties and Methodological Approaches 250 Operational Categories of Single-Cell Activity 259 Regulation of Physiological Structure of Marine Bacterioplankton 260 Factors Influencing Physiological State of Bacterial Cells in Marine Ecosystems 261 Factors Influencing Loss and Persistence of Physiological Fractions 263 Distribution of Single-Cell Characteristics in Marine Bacterioplankton Assemblages 265 Distribution of Single-Cell Activity and Physiological States in Marine Bacterioplankton 265 Simultaneous Determination of Several Aspects of Single-Cell Activity and Physiology 270 Patterns in Distribution of Single-Cell Activity and Physiology Along Marine Gradients 271 Distribution of Activity and Growth Among Bacterial Size Classes 273 Distribution of Activity Across and Within Major Phylogenetic Groups 274 Dynamics of Single-Cell Activity and Physiological States 276 Ecological Implications of Patterns in Bacterioplankton Single-Cell Activity 279 Community Versus Individual Cell Growth and Metabolic Rates 280 Linking the Distribution of Single-Cell Parameters and the Bulk Assemblage Response 282 Ecological Role of Different Physiological Fractions 283 Concluding Remarks 284 Summary 285 Acknowledgments 285 References 285 9 HETEROTROPHIC BACTERIAL RESPIRATION 299 Carol Robinson Introduction 299 Measurement of Bacterial Respiration and Production 301 Routine Measurement Techniques for Bacterial Respiration and Their Limitations 301 Routine Measurement Techniques for Bacterial Production and Their Limitations 304 x CONTENTS
- 15. Magnitude and Variabilityof Bacterial Respiration 304 Temporal Variability 308 Spatial Variability 309 Relationship Between Bacterial Respiration and Environmental and Ecological Factors 311 Bacterial Respiration as a Proportion of Community Respiration 315 Predicting Bacterial Respiration 317 Comparison Between Measurements and Predictions of Bacterial Respiration 319 Magnitude of Bacterial Respiration in Relation to Primary Production 321 Bacterial Respiration in a Changing Environment 324 Summary 326 Acknowledgments 327 References 327 10 RESOURCE CONTROL OF BACTERIAL DYNAMICS IN THE SEA 335 Matthew J. Church Introduction 335 Growth in the Sea 336 Growth and Nutrient Uptake Kinetics 339 Approaches to Understanding Resource Control of Growth 343 Comparative Approaches 343 Experimental Approaches for Defining Limitation of Bacterial Growth 349 Limitation by Dissolved Organic Matter 351 Bacterial Growth on Bulk DOM Pools 353 Limitation by Specific DOM Compounds 354 Limitation by Inorganic Nutrients 361 Nitrogen 361 Phosphorus 364 Trace Nutrients 365 Temperature–DOM Interactions 366 Light 368 Resource Control of Specific Bacterial Populations in the Sea 369 Summary 371 Acknowledgments 371 References 371 CONTENTS xi
- 16. 11 PROTISTAN GRAZINGON MARINE BACTERIOPLANKTON 383 Klaus Jürgens and Ramon Massana Introduction 383 New Insights into Phylogenetic Organization 386 Functional Size Classes of Protists 390 Natural Assemblages of Marine Heterotrophic Nanoflagellates 391 Functional Ecology of Bacterivorous Flagellates 394 Living in a Dilute Environment 394 Using Culture Experiments to Infer the Ecological Role of HNF 397 Impact of Protistan Bacterivory on Marine Bacterioplankton 401 Search for the Perfect Method to Quantify Protistan Bacterivory 401 Rates of Protistan Bacterivory in the Sea 403 Balance of Bacterial Production and Protistan Grazing 404 Bottom-Up Versus Top-Down Control of Bacteria and Bacterivorous Protists 405 Ecological Functions of Bacterial Grazers 406 Grazing as a Shaping Force of Bacterial Assemblages 408 Bacterial Cell Size Determines Vulnerability Towards Grazers 408 Other Antipredator Traits of Prokaryotes 411 Grazing-Mediated Changes in Bacterial Community Composition 413 Molecular Tools for Protistan Ecology 414 Culturing Bias and Molecular Approaches 414 Global Distribution and Diversity of Marine Protists 420 Linking Diversity and Function for Uncultured Heterotrophic Flagellates 422 Summary 423 Acknowledgments 424 References 424 12 MARINE VIRUSES: COMMUNITY DYNAMICS, DIVERSITY AND IMPACT ON MICROBIAL PROCESSES 443 Mya Breitbart, Mathias Middelboe, and Forest Rohwer Introduction 443 Viruses and the Marine Microbial Food Web 444 Direct Counts and Viral Numbers 444 xii CONTENTS
- 17. Viral Production andDecay Rates 447 Viral Decay and Rates of Production in Pelagic Systems 447 Measurements of Viral Production in Marine Sediments 449 General Rates of Viral Production 449 Role of Viruses in Biogeochemical Cycling 450 Impact of Viruses on Bacterial Diversity and Community Dynamics 452 Marine Viral Diversity 457 Methods for Examining Marine Viral Diversity 457 Culture-Based Studies of Viral Diversity 458 The Need for Culture-Independent Methods 459 Culture-Independent Studies of Viral Diversity Using Transmission Electron Microscopy 460 Whole-Genome Profiling of Viral Communities Based on Genome Size 461 Studies of Viral Diversity Using Signature Genes 461 Metagenomic Studies of Viral Diversity 462 A Vision for the Future 466 Summary 467 References 468 13 MOLECULAR ECOLOGICAL ASPECTS OF NITROGEN FIXATION IN THE MARINE ENVIRONMENT 481 Jonathan P. Zehr and Hans W. Paerl Introduction 481 Chemistry, Biochemistry, and Genetics of N2 Fixation 482 Genetics and Enzymology 483 Evolution of N2 Fixation 485 Phylogeny of Nitrogenase 487 Genomics of N2 Fixation 487 Diversity of N2-Fixing Microorganisms 489 Regulation in Diazotrophs 489 Methods for Assessing Diazotroph Diversity, Gene Expression, and N2 Fixation Activity 490 Ecophysiological Aspects of N2 Fixation 494 Ecology of Diazotrophs in the Open Ocean 499 Estuarine and Coastal Waters 505 Benthic Habitats, Including Microbial Mats and Reefs 506 Deep Water and Hydrothermal Vents 507 Summary 508 Acknowledgments 509 References 509 CONTENTS xiii
- 18. 14 NITROGEN CYCLINGIN SEDIMENTS 527 Bo Thamdrup and Tage Dalsgaard Introduction 527 Inputs 531 Transformations 532 Microbes and Microbial Processes 532 Processes Involving Mn and Fe 548 Nitrogen Budgets 550 Benthic Budgets 550 Oceanic Budgets 552 Summary 554 References 555 INDEX 569 xiv CONTENTS
- 19. PREFACE It has beennearly 10 years since work started on the first edition of this book. Ten years is a long time for just about any field of science, but especially for a fast-moving one such as marine microbial ecology. Here, finally, is the second edition. This book is more than just a revision of the first edition which was published back in 2000. Some chapters from that edition are not repeated here, because work in those areas has slowed and the basic principles covered before have not changed. However, those topics and principles remain as important and as valid today as 10 years ago, and readers should hang onto the 2000 book (or get it if they do not have it already); much of it is still relevant. Other chapters of this book have titles similar to those in the first edition, but even in these cases, the chapters have been substan- tially rewritten, often by authors who have different perspectives on the topics covered previously. Finally, several chapters discuss microbes and biogeochemical processes that we were just beginning to learn about 10 years ago, and still others that we did not even know existed back then. What remains the same is the intended audience: advanced undergraduates, begin- ning graduate students, and colleagues from other fields wishing to learn about microbes and the processes they mediate in marine systems. This book, aided by the first edition, is meant to be as close to a textbook as a multi-authored book can be. I wish to thank several people who helped to get this book published. First and most importantly, I thank the chapter authors for agreeing to work on this project and also for looking over another chapter (or two) in the book. Each chapter was reviewed by another chapter author and an outsider not connected to the book. I especially want to thank Jens Boenigk, Hugh Ducklow, Pete Conway, Stefan Hulth, Rick Keil, Karin Lochte, Alison Murray, Jack Middelburg, Jarone Pinhassi, Thomas Reinthaler, Janice Thompson, Daniel Vaulot, Tracy Villareal, and Peter Williams. The anonymous reviewers (and those I’ve forgotten to mention by name—sorry) also deserve thanks. I greatly appreciated Dave Karl’s support during a critical junction of this project, and I acknowledge the help of Karen xv
- 20. Chambers, Thom Moore,and others at Wiley. Finally, I thank the readers of the first edition of this book. This second edition would not have come about if not for your positive comments and feedback. DAVID KIRCHMAN Lewes, Delaware xvi PREFACE
- 21. CONTRIBUTORS ODED BÉJÀ Facultyof Biology, Technion–Israel Institute of Technology, Haifa, Israel [[email protected]] MYA BREITBART College of Marine Science, University of South Florida, St. Petersburg, FL 33701, U.S.A. [[email protected]] MATTHEW J. CHURCH Department of Oceanography, University of Hawaii, Honolulu, HI 96822, U.S.A. [[email protected]] TAGE DALSGAARD National Environmental Research Institute, University of Aarhus, DK-8600 Silkeborg, Denmark [[email protected]] JED A. FUHRMAN Department of Biological Sciences, University of Southern California, Los Angeles, CA 90089, U.S.A. [[email protected]] JOSEP M. GASOL Institut de Ciències del Mar, CMIMA (CSIC), Passeig Marı́tim de la Barceloneta 37–49, 08003 Barcelona, Catalunya, Spain [[email protected]] PAUL A. DEL GIORGIO Département des Sciences Biologiques, Université du Québec à Montréal, CP 8888, succ. Centre Ville, Montréal, Québec, Canada H3C 3P8 [[email protected]] ÅKE HAGSTRÖM Kalmar University, Sweden [[email protected]] KLAUS JÜRGENS Leibniz Institute for Baltic Sea Research, 18119 Rostock, Germany [[email protected]] DAVID L. KIRCHMAN College of Marine and Earth Studies, University of Delaware, Lewes, DE 19958, U.S.A. [[email protected]] RAMON MASSANA Institut de Ciències del Mar, CMIMA (CSIC), Passeig Marı́tim de la Barceloneta 37–49, 08003 Barcelona, Catalunya, Spain [[email protected]] MATHIAS MIDDELBOE Marine Biological Laboratory, University of Copenhagen, DK-3000 Helsingør, Denmark [[email protected]] xvii
- 22. MARY ANN MORANDepartment of Marine Sciences, University of Georgia, Athens, GA 30602-3636, U.S.A. [[email protected]] TOSHI NAGATA Ocean Research Institute, The University of Tokyo, Tokyo, Japan [[email protected]] FABRICE NOT Evolution du Plancton et PaleOceans Laboratory, CNRS, Université Paris 06, UMR7144, Station Biologique de Roscoff, 29682, Roscoff, Cedex BP 74, France [[email protected]] HANS W. PAERL Institute of Marine Sciences, University of North Carolina at Chapel Hill, Morehead City, NC 28557, U.S.A. [[email protected]] CAROL ROBINSON School of Environmental Sciences, University of East Anglia, Norwich NR4 7TJ, U.K. [[email protected]] FOREST ROHWER Department of Biology, San Diego State University, San Diego, CA 92182, U.S.A. [[email protected]] EVELYN SHERR College of Oceanic and Atmospheric Sciences, Oregon State University, Corvallis, OR 97331-5503, U.S.A. [[email protected]] BARRY SHERR College of Oceanic and Atmospheric Sciences, Oregon State University, Corvallis, OR 97331-5503, U.S.A. [[email protected]] MARCELINO T. SUZUKI Chesapeake Biological Laboratory, University of Maryland, Center for Environmental Science, Solomons, MD 20688, U.S.A. [[email protected]] BO THAMDRUP Institute of Biology, University of Southern Denmark, DK-5230 Odense M, Denmark [[email protected]] ALEXANDRA Z. WORDEN Monterey Bay Aquarium Research Institute, Moss Landing, CA 95039, U.S.A. [[email protected]] JONATHAN P. ZEHR Ocean Sciences Department, University of California, Santa Cruz, CA 95064, U.S.A. [[email protected]] xviii CONTRIBUTORS
- 23. Figure 6.8 Epifluorescenceimages of picoeukaryotes from environmental samples illumi- nated via FISH. (a, b) Surface samples from the English Channel hybridized with a Chlorophyta-specific probe (CHLO02) and a Micromonas genus-specific probe (MICRO01), respectively, using TSA-FISH. The cytoplasm of hybridized cells has green fluorescence (FITC dye). (c, d) Florida Straits samples from 5 m and 85 m depth, respectively, hybridized with the Haptophyta-specific probe (PRYM02) using the TSA-FISH technique and FITC. The natural red/orange fluorescence of the cyanobacteria can be seen. The sample shown in (d) has been counterstained with the DNA-specific dye DAPI (blue) to allow visualization of hetero- trophic bacteria. (e, f ) A surface sample from the Blanes Bay (Mediterranean Sea) hybridized with the probe NS4 (Massana et al. 2004), specific for MAST-4 cells. In contrast to other images shown, the FISH method used to illuminate MAST cells in ( f ) did not involve signal amplification, but rather used a monolabeled probe linked to the fluorescent dye CY3 (red). The DAPI counterstaining from the same microscopic field is shown in (e). Scale bars represent 10 mm, except in (e, f ), for which they represent 20 mm. (a) Courtesy of F. Not. (b) Courtesy of E. Foulon. (c, d) Courtesy of J. A. Hilton, F. Not, and A. Z. Worden. (e, f ) Courtesy of R. Massana.
- 24. Figure 13.4 Photomicrographsof cultivated and uncultivated marine cyanobacterial diazo- trophs. (a) Uncultivated unicellular cyanobacteria from the open ocean that are likely diazo- trophs. (b) Diversity of Trichodesmium morphology in natural populations. (c, d) Symbiotic unicellular cyanobacteria. (e) Heterocystous symbiont of diatoms. ( f ) Heterocystous symbiont of diatoms. (g) Heterocystous symbiont of diatoms. (h, i) Association of bacteria with hetero- cysts of heterocyst-forming cyanobacteria.
- 25. Figure 13.6 Remotesensing of blooms of Trichodesmium. (a) True-color image of surface ocean, indicating position of Trichodesmium bloom detected by remote sensing. (b) Remote sensing image of Trichodesmium using the algorithm of Subramaniam et al. (2002). Modified from Subramaniam et al. (2002) and used with permission from Elsevier.
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- 27. It was withno small satisfaction I arrived at the now familiar homestead of Núpstað, and received the usual glad welcome from the bóndi Ayólver, who had been expecting us. I again took up my quarters in the disused little church, which makes such a good storehouse for my friend Ayólver, and such an excellent resting-place for chance travellers like myself. It seemed quite home-like as I tumbled into the little bed which had been improvised upon the boxes in the corner, and I experienced the comfortable feeling of being in my old place again as I ate my breakfast off and posted up my diary upon the antiquated communion table. Do not be shocked, good reader! all sanctity had long ago departed from this useful piece of furniture, and if we were to peep into the inside, we should find neither sacred utensils nor vestments; but simply the serviceable homespun garments of the bóndi’s wife. The farm and the rocks behind it were but little altered since I first saw them four years ago. One year in Núpstað is much like its predecessor, and things go on, year after year, in just the same routine, except where the inevitable changes of life and death intervene. The people had altered the most, for of course they had grown older, and one or two faces were missing! Well, I have grown older, too—it is no good to stand dreaming. There is a bullock to be bought, butchered, and salted, preparatory to making it into “kœfar,” as the Icelanders call the kind of pemmican I make for my Jökull expeditions. Skin-bags and mocassins have to be procured; butter, bread, and stock-fish have to be sought after; in fact, the greater part of three weeks’ provisions for six men must be collected from the neighbouring farms. We made the necessary arrangements, and settled that these various articles are to be ready for us in a week’s time; we then deputed Paul’s father to attend to the levying of our requisitions, and the payment for them. The ox was next slain, dissected, and salted, and we were again ready to start on our travels. Some little difficulty was experienced in getting all into train, owing to the hurry all the farmers of this locality were in to get this year’s wool to the store at Papós, which is situated four days’
- 28. journey to theeast; for tidings had been received that the ice of a portion of the Vatna Jökull, known as the Breiðamerkr had advanced to such an extent as to threaten the cutting off of all communication along the sea-shore, since the advance still continued. In consequence of this alarm every farmer was busy preparing the wool for market; steaming cauldrons were cleansing it from its grease, bands of sturdy Icelandic maidens were rinsing it in the clear water of the mountain streams—which are almost sure to be found in close proximity to the farms in this part of the country—patches of white wool were drying upon the ground, while the male part of the community were measuring it in quaint wooden baskets, packing it into sacks, and forming bundles of equal weight to balance on each side of the pack-horses. It would be a very serious thing, indeed, if the road to Papós were to be intercepted, as it would compel the dwellers in this district to journey to Eyrarbakki before they could exchange their produce for the necessaries they require. Leaving Núpstað behind us, we set out for the advancing glacier, and turned our faces towards the snowy slopes of Örœfa. The Súla river, or Núpsvatn, had to be crossed. It was deeper than I had before seen it, though its volume of water scarcely seemed to have increased. Its bed was changed to one of pebbles and quicksand. In 1871 it was of pebbles only, in 1874 it was black sand, in 1875 it is again pebbles and sand. We crossed the river and fast sped on our way over the desert of Skeiðarár Sandr. This sand occupies an area of 300 square miles. It has been formed by the joint efforts of volcanoes upon the Vatna and Mount Örœfa, which have strewn this tract with sand and ashes, and whose ejectamenta have been brought down by the shifting waters of numerous glacial streams which traverse the Skeiðarár Sandr in many directions. It would seem that the portion of the Vatna which here bounds the Skeiðarár Sandr upon the north has acted in a similar manner to the Breiðamerkr Jökull; for numerous moraines occur upon these sands, some of which are at a great distance from the utmost limit of the Jökull at the present time. Indeed, there has been an obvious advance at this point since 1871
- 29. of the fringeof the glacier which almost surrounds the Vatna Jökull. The existence of scratched rocks in moraines in Iceland below the limit of the glaciers does not of necessity prove that such glaciers have bodily advanced, as during extensive eruptions of glacial mountains huge masses of ice frequently slip forward to considerable distances, scratching the harder and furrowing the softer rocks in their progress, which, upon their melting, leave large piles of glacial débris, in no way distinguishable from a moraine stranded upon the lower elevations. It was blowing hard from the east with heavy rain, but upon the west side of the mountain before us (Örœfa) the sun was shining in the most tantalising manner, so that as we urged our horses along the heavy sands we were fain to fancy ourselves exploring those dazzling glaciers and snowy slopes which seemed to fascinate the sunshine and detain it from reaching us. We were soon under the lee of the mountains before us. Sheltered from the wind and the storm, we could stop to admire the grand sweep from the Örœfa to the commencement of the Skeiðará Jökull. Looking back at Núpstað, we saw it enwrapped in gloom, the clouds clustering round the Lómagnúpar,[1] a mountain which seems to attract all the bad weather to Núpstað and the storm sat heavily upon the western portion of the plain of Skeiðarár Sandr, which was exposed to the fury of the east winds. Crossing the river Skeiðará, we reached the Saga-famed Svínafell. Here we stayed to refresh ourselves with the national panacea for the ills of Icelandic travel, namely, a cup of coffee of the real Icelandic brew! The art of making good coffee is one of the greatest accomplishments of the fair sex here, and it is a pity it is not more generally attained by the lady population of other countries. The occurrence of drinkable coffee in Iceland, a good cup of it being always to be obtained at the poorest farm, is the more remarkable, as the coffee sold by the merchants at the various stores is never of the best quality; but is principally the Java coffee. The grand secret of success in this special domestic art is doubtless owing to the fact
- 30. that the coffeeis roasted at home, exactly to the right turn, and deftly manipulated in some particular way which early training and long practice can alone effect. The last and by no means the least adjunct to this national bonne bouche is in most cases a good supply of cream. Being thus fortified, we were taken to see a birch-tree upon the hill behind the farm. This tree might have been five-and-twenty feet in height, and it was considered the largest tree in this part of the island. There is, however, a considerable growth of bushy trees, principally birch, in the valley called Núpstaða-skógr down which the river Súla flows. It is by far the largest wood in the south of Iceland. Núpstaða-skógr is likewise remarkable for containing a breed of wild sheep, which belongs to our friend Ayólver, who is the owner of the skógar, together with the valuable farm of Núpstað. There is also another patch of wood at the north-west base of Örœfa, which is of great use to Svínafell and the adjacent farms. The hills behind Svínafell are basaltic; but as we proceeded further eastward, we soon found ourselves surrounded by the more recent products of the volcano Örœfa, which towered above us upon our left hand. Seeing a party of horsemen approaching, we whipped our little drove together, and met them upon the grass which was a few hundred yards off. The party consisted of an Althing’s-man, who was going to Reykjavík to attend the Althing, or Icelandic Parliament, with his servants, and the priest from Sandfell, at whose house he had been staying, and who was escorting him for a short distance. The priest turned out to be a cousin of my man Paul, so after a brief colloquy, and requesting the Althing’s-man to convey our greetings to friends at Reykjavík, we rode on to Sandfell. Our road lay past several beds of white pumice which had all been ejected from Örœfa. A smart gallop over cinders and fragments of lava brought us to the church and parsonage. Sandfell is situated at the south base of Örœfa. Behind it rise barren hills of compact agglomerate, composed of volcanic ash and fragments of lava, but
- 31. our friend thepriest is compensated for his dreary surroundings by having one of the prettiest Icelandic women I have seen for his wife. She seemed quite piqued because I could not own to thinking Sandfell a very pretty place. Going hence, we crossed the stream of lava and agglomerate, which I was informed resulted from the eruption of Örœfa in 1862. This stream is a remarkable one, inasmuch as the agglomerate has flowed down in a semi-molten state, cotemporaneously with the lava, both being mixed together; the agglomerate appears to preponderate, but this may be the result of the lava being of higher specific gravity, which causes it to sink to the bottom of the stream. We stopped for the night at Myrum,[2] on the south-west of the Breiðamerkr Sandr. The bóndi, like all the people of this district, was hastening to get to Papós with his wool. We supped and breakfasted off some birds which our host called Svartfugl. They were the nicest birds I had ever tasted in Iceland, the meat being tender and plenty of it, and I thought so well of this dish that I took one of the birds away with us for our lunch on the road. Here we hired a fresh horse, leaving Paul’s, which had contracted a sore back, and started over the Breiðamerkr Sandr. The sands, like the Skeiðarár Sandr are the result of the great efforts of the Örœfa and Vatna Jökulls, more especially the part of the Vatna known as the Breiðamerkr Jökull, which was the one whose movements we had to examine. The road over these sands is long and dreary, especially in such weather as had just overtaken us. We passed an extensive encampment of farmers, who were on their way to Papós; but, despairing of crossing the rivers which traverse the Breiðamerkr Sandr upon such a day with heavily laden horses, they had decided on remaining encamped upon the little patch of grass they had reached. About one third of the way over the Sandr we arrived at the farm of Kvísker, which is situated upon a little oasis of grass- land. We found it a very acceptable halting-place, and although we were wet, we were glad to sit down and take coffee and schnapps,
- 32. and smoke apipe inside; the room had no windows, and it was filled with planks and carpenter’s tools, for the house was being enlarged. We could obtain but little food for our horses, and the greater part of our day’s work had yet to be accomplished, so a quarter of an hour saw us again to horse, and rapidly approaching the extreme point of the advancing Jökull. This Jökull appeared unlike most of the Icelandic glaciers I have seen. Instead of terminating in an even slope, or steep rounded cliffs of ice, sometimes fissured, but generally very regular, it terminated in an irregular wall of cloven and contorted masses of the rifled and dislocated glaciers; while the more elevated masses assumed the form of spires, towers and grotesque architectural shapes. As we were intently looking at them, some of them tottered and fell. It is indeed a serious matter to contemplate the short distance now left between the Jökull and the sea—at one point not more than 250 yards—in addition to this, new rivers have been formed between the Jökull and the sea, which have to be crossed, but which it would be impossible to do with a strong south wind blowing. The Jökulsá is quite bad enough, but to have several miles of road converted into quicksand by the diverted waters of the Jökulsá, and to have new rivers in addition to the advance of the Jökull, is enough to make the people of the district fear for the road to Papós. One consolation may exist—that the Jökull has advanced before, and, after a considerable time, retreated. Still, as an old inhabitant of the neighbourhood informed me, “It never has advanced as it does now,” and even upon the other occasion, upon the whole, it gained ground. Alas! poor Iceland —both fire and water appear allied against it; the latter especially, in all its forms—boiling, cold, and frozen, and in the form of rain, hail, snow, and vapour! We were obliged at one point to travel along the sea-shore, where we espied the body of a large fish with some dark objects moving about it. A nearer approach showed it to be a small whale, which, from olfactory evidence, had lain there for some time. The dark objects, startled at our appearance, rose in a covey of— well, the same birds of which we had enjoyed the flavour at Myrum. Svartfugl have never tasted quite so nice to me since. At last the Breiðamerkr Sandr were passed; fresh mountains rose before us,
- 33. and the weathercleared. To our right was a remarkable lagoon, Breiða-bólstaðalón; which is a narrow fjord, twelve miles in length, enclosed upon the south by a large sand-bank running parallel with the shore. This lagoon is open to the sea at the north-east end, but is too shallow for ships to enter. Evening found us at Kálfafellstaðr, a place pleasantly situated beneath the outlying hills of the Vatna Jökull. These hills are principally composed of amygdaloidal basalt, abounding in zeolites; chalcedonies are especially plentiful, and I dare say it might pay to look for the precious opal. This eastern corner of Iceland appears to be particularly rich in zeolites; I noticed the same when I was at Berufjörðr. We stayed for the night with another relative of Paul—he seemed to have kindred nearly all over the island, and a very superior race they appear to be. This relation was the widow of the former priest of Kálfafellstaðr. Here we bought another horse, and hired the widow’s son, a lad about seventeen; for we required a man and a lad to drive our horses round to the north of the island while we crossed the Vatna Jökull. The widow and her daughter accompanied us a short distance upon our return journey, and, after two days’ riding, we were again at Núpstað. Preparations for our journey across the Vatna now commenced in earnest. The sleighs and the snow-shoes had been made according to our instructions. All was there except the men and the butter; enough of the latter, however, turned up in the morning to enable us to make the pemmican, which I at once set myself to work to superintend. A fire was lighted and a cauldron of water soon heated, and the beef boiled; then came the work of cutting up an entire ox into pieces the size of ordinary wine-corks. Paul senior, and I commenced operations by first taking out the bones; and, by dint of sharp knives, and a few hours’ hard work, we prepared about seventy- eight pounds of meat. Twenty pounds of salt butter and half-a- pound of salt were then melted in the cauldron, and the meat
- 34. carefully mixed withit. After a short time it was ready to be packed in the skin bags in which it was to be carried. The bags were placed in troughs of water during the operation of filling, to prevent leakage at the seams, and when they were filled they were tied up and laid in a stream close by, where stones were piled upon them to press down the meat. When they were sufficiently pressed, and the contents had become cold (which took about twenty hours), they were each placed in ordinary sacks for more easy carriage; for greasy skin-bags full of meat are rather slippery things to carry, and somewhat nasty things to handle. By June 25th all my preparations were made, and my men arrived; Paul Paulsen and a cousin of his from Skaptarfellssysla; Helgi, from the farm of Króki; Finnur, from Myrdalssysla; and Eyólfur, from Hörgsdalr: these were to accompany me across the Vatna Jökull. In addition were Bjarni, who was with me last year; the farmer from Rauðberg, who carried the post between Prestbakki and Berufjörðr—a deaf and dumb man, and a man named Vigfúss; these four were to return when we reached the mountain which I last year named “Mount Paul,” about a third of the way across the Jökull. I had also arranged with Paul’s father and little Arni, whom I had hired at Kálfafellstaðr, to take our horses from Núpstað round the east side of the Vatna into the north of the island. Our equipment, which was to be drawn upon hand-sleighs, consisted of a low tent, four feet high; a large sleeping-bag, which would accommodate six of us—this was eight feet long, and five feet wide—one side being made of a layer of cork and felt, covered with mackintosh, and the other of a stout blanket also covered with waterproof. This bag was open at both ends, so that three could sleep with their heads one way and three with their heads the other. Both these openings were covered by a hood, which proved a great protection to our heads while sleeping, and prevented the snow from getting into the bag. This gave us sleeping accommodation for six persons, with a weight of only sixty pounds. This bed, however, had its disadvantages; for instance, if any one was taken with cramp, or
- 35. dreamt of engagingin any particularly active exercise, its limited dimensions became painfully apparent; moreover, it is almost impossible to keep the inside of the bag perfectly dry, owing to the exhalation from our bodies. I have paid great attention to this matter, but have found that for a prolonged sojourn amidst wet snow, where weight is a subject of paramount importance, it is the best sleeping arrangement that can be contrived. Our provisions consisted of 100 lbs. of pemmican in skin bags, 50 lbs. of butter, 100 lbs. of skonrok, or Danish ship-biscuits, 15 lbs. of dried fish, 15 lbs. of dried mutton, 15 lbs. of gravy soup, 2 tins of “soupe Julienne,” in packets; 6 tins of chocolate and milk, 2 lbs. of cocoa, and 4 lbs. of sugar; 2 gallons of proof whiskey, 1 gallon of spirit for burning, 5 lbs. of tobacco, and 3 tins of Peek and Frean’s meat biscuits. I had a small Russian furnace, which is an excellent lamp for heating water or melting snow. These articles, with a good supply of warm clothing, waterproofs, and mocassins (for it is impossible to wear leather boots in the snow), and the necessary instruments and implements, completed our outfit. All things were now ready, and the day had at length arrived when we were to assail the Vatna again. We rose betimes, but it was midday before we were fairly on our way. I took leave of the bóndi Ayólver, who would not charge me anything for my own board and for the keep of my own horses. He was too unwell to accompany us to the Vatna, and seemed quite upset at saying good-bye, as he said he felt sure it would be for the last time, whether we got across the Jökull or not. I cheered him up, and said, I hoped some day or another to come to Núpstað again; and so we started on horseback, and, after crossing the river Diúpá, we commenced the ascent of Kálfafellsfjall, which hill lay between us and the Vatna. The journey was a very trying one to the horses; it is so at the best of times, but now the melting snow still lay thickly, and in places had converted the unstable soil into quicksands. In some parts it was necessary to cross ravines full of snow, which had melted underneath, leaving the bottom of the ravine roofed. The
- 36. horses fought veryshy of these snow-roofed valleys, and when we came to any hole which had been formed by the subsidence of a portion of the snow into the valley beneath, it was with difficulty we could get them along, as the noise of the stream, which invariably ran below, made them rather fractious. But the snow having regelated into an indurated compact mass, was often some yards in thickness, so I do not think there was any real danger of sinking through it. These preliminary difficulties were soon disposed of, and 6 P.M. found us at that point where the rocks terminate and the eternal snows of the Vatna commence. A squall of sleet and wind now rolled down upon us. I immediately directed two men to prepare some coffee, for we had brought wood for that purpose, while some gave the horses a feed of hay, and others unpacked the burdens they had carried so pluckily from Núpstað. The coffee was soon ready, the storm cleared, and the scene must have bordered on the picturesque, or perhaps the “unique,” as we all clustered round the remnant of the fire, amid the different packages that were to cross the Vatna, our horses pawing the ground, impatient to return to their pastures. The grand white Jökull lay before us, the black crags of the fjalls behind us, and the roar of the Diúpá in our ears, while a beautiful rainbow spanned the eastern sky—a harbinger, we trusted, of good success. Here we took leave of Paul’s father and his cousin Arni, directing them where to wait for us with the horses, in the north of the island. The evening promised to be showery; but having a lively reminiscence of the black sand of this locality, which at our last year’s encampment upon this spot got into our ears, our eyes, and our food, I determined to advance and camp, as soon as we needed to do so, upon the deep snow, although my men had already begun to put up a temporary abode with loose stones from the terminal moraine of the Jökull. At this point last year the Jökull was a crevassed glacier, whose surface was covered with aiguilles and hummocks of black sand and ice. But all traces of the glacier were buried beneath a vast
- 37. accumulation of snow!From the first we were able to use our sleighs, and, turning due northward, we left the habitable world behind us, being face to face with the hardest piece of our summer work. As far as the eye could see was one lifeless, pathless wilderness of snow, destitute alike of animal, insect, or floral life. Our footsteps gave no sound, and our very voices seemed strange in this drear solitude, the death-like stillness of whose snowy wastes is broken only by the howling of the storm, or the outburst of a volcano! It was evident that a much greater snowfall had taken place during the past winter than in the preceding one, and the newly-fallen snow took us up to our knees, making our progress very difficult and slow. After about three hours’ dragging, it began to snow, and a thick fog enveloped us, so I decided to encamp. The plan I usually adopt for sleeping in the snow—and I believe one of the warmest and best methods—is to dig a square hole, three or four feet deep; over this I pitch my tent, banking it well round the sides with snow. I then spread the sleeping bag at the bottom of the hole, with the hoods doubled down over the ends to prevent any snow getting into it. If a storm is blowing, I cast up a bank of snow to windward, and take everything that will be required for immediate use into the tent. The next thing is to draw the sleighs up to the door of the tent; so that if anything extra is required it can be procured without much difficulty, and having stuck up all sticks and shovels firmly in the snow, to prevent their getting covered up and lost, we turn in, changing our wet or snowy clothes sitting upon the waterproof exterior of the bag, and, putting on a dry change, we all get into the bag, having previously fixed up waterproof coats upon the snowy wall at each end, to lean against. If it is not freezing very hard, we hang our snowy clothes upon a line at the top of our tent, with our satchels, &c.; but if it is freezing hard we put them underneath the bed. Snow is then melted, soup or chocolate is made, and rations served, which, with a small allowance of grog, pipes, and a song all round, finish our labours for the day or night, as the case may be, and we go to sleep.
- 38. This was themanner in which we now camped, six of us occupying the sleeping-bag, much after the manner of sardines in a sardine box, the remaining four, who were only to accompany us as far as Mount Paul, made themselves as comfortable as they could with rugs and mackintosh coats in the front part of the tent. I ordered every man to fill his flask with snow and put it in his pocket, that each might have a drink of water when he awoke, and in the course of an hour nothing could be heard but the heavy, stentorian breathing of nine out of ten of our party. Having posted up my diary, I slept well for an hour, when I was awakened by a sudden commotion at the other end of the tent. I called out to Paul for an explanation, saying, “Holloa! what’s the matter at your end?” He replied in a deep, solemn voice, “Now is the dumb beating his feet.” Although our dumb friend’s feet were doubtless cold, I could not allow that method of warming them in a tent only 10 by 6½ feet, and I therefore directed that another man should chafe the dumb man’s feet and cuddle them up in his arms. The morning brought us only fog and storm, but after a few hours the latter abated. I served out some warm soup, and we got under weigh. After an hour the fog became so dense, the snow so soft and deep, and a determined sleet had set in, that I was obliged reluctantly to call a halt. Between nine and ten in the evening the weather cleared, the wind shifted to the north-west and the sun came out, and we again advanced; but the snow being up to our knees, I perceived I was tiring my men. So after going on a few miles I again halted, as it had begun to freeze, and the probability was that in about two hours the snow would be firm enough to travel on. Casting up a bank of snow to windward, we six turned into our bag upon the surface of the snow, leaving the tent and all other wraps for our four extra men. It was bitterly cold, but the atmosphere was very clear. By 3 A.M. I roused my men; the thermometer registered 20° Fahrenheit; a firm crust had formed upon the snow which bore us bravely. It was a glorious morning and a stiff north wind was blowing; the sleigh travelled merrily along, and as the sun illumined the magnificent snow slopes around us, everything seemed to promise fine weather
- 39. and success. Thepure element we were breathing seemed to give us fresh life and strength, and made us feel equal to the work before us. After three hours one of the men (Vikfúss) gave out, said he could go no further, and lay down upon the snow; but as there were not nearly so many degrees of frost now, the man was warmly clad, and I had a great idea he was shirking, I left him behind, much against the will of his companions. Before we were half a mile away I had the satisfaction of seeing him following, apparently not very much the worse for wear. The ascent from the first had been a very gradual slope of snow, which now became undulating and somewhat steeper, especially upon the N.E., where steeps of snow swept up to the mountain. I last year named Vatna Jökull “Housie,” from the great resemblance which its summit, then free from snow, bore, when viewed in one aspect, to the roof of a house. The likeness was now much less striking, from its being all white. I can scarcely go on without remarking upon the excellence of the postman from Rauðberg. He was always cheerful, willing and obliging, and had twice the hardihood and strength of the other men. I only regretted I could not take him right across the Vatna, but his postal duties would not admit of so prolonged an absence. We sighted Mount Paul at 9 A.M. Here we made a good breakfast, and our disabled man having slunk up, he made better progress with his meal than he did with his sleigh. Mount Paul is a cluster of one large and several smaller volcanic eminences, rising to the height of 150 feet above the surrounding snow. A semi-circular pit being thawed out by the radiation of the sun’s rays from the south side of the mountain, we found here an abundant supply of water. The mountain is composed of varieties of obsidian, varying from the highly vitreous to the grey stony variety; one portion of it consists of vitreous obsidian cementing together multitudes of the concretionary forms commonly known as spherulites. We slept for two or three hours; but the state of the snow was such that it was impossible to get the sleighs through it. I sent back
- 40. my four extramen, for they had little or nothing to carry, and we had left them a good supply of provisions at the commencement of the Jökull. As the accommodation in the tent was but small for them, and it seemed to promise bad weather, they preferred forcing their way back through the soft snow to running the chance of being weather-bound for three or four days. They had not been gone away many hours when it began to rain, and as night drew on it became more and more evident that there would be no frost. The wind had shifted to the S.S.E., the thermometer stood at 33° Fahr., and as the night advanced the snow became so soft and rotten that in some places it took us up over our knees. The next day the wind was still S.S.E., and the fog and sleet were as bad as ever; and as progress was impossible, I minutely inspected the rocks of Mount Paul. They rise from a large crater now filled with snow. To the south-east is a pit-crater partially filled with snow. Mount Paul is composed almost entirely of perlite and obsidian. This is the only place in Iceland in which I have found obsidian “in situ.” The west side of the mountain particularly attracted my attention, being composed of multitudes of spherulites cemented together by obsidian. Thousands of these small globular formations had been weathered out of the obsidian, and in some places one might have collected a hat-full. Night brought no improvement in the weather; and a somewhat remarkable scene presented itself of six men lying in a hole in the snow, 4250 feet above the sea-level, in Iceland, all hoping for a frost —but no frost came, and morning found us in the same position. This was very aggravating for one who had spent much money, time and labour, in order to complete a survey across the Vatna Jökull; but the day was fine, and I could post up my diary, plan for the future, learn Icelandic, eat, drink and smoke, upon the volcanic débris on the leeward side of Mount Paul, where the thermometer at midday rose to 75 and 80 degrees in the sun, and it was infinitely preferable to lying in the snow. Towards evening it began to freeze, so we packed up our sleighs and retired to Mount Paul, until the crust was strong enough to bear the weight of the sleigh. By ten P.M.
- 41. there were twelvedegrees of frost, and the wind blew freshly from the N.W. The crust now bore the sleigh, but we sank through it up to our knees at every step. This was such laborious work that after two hours we halted, hoping the crust would soon become firmer; but we were doomed to disappointment, for after a while the wind suddenly shifted to the S.E., and almost simultaneously a fog appeared. However, we were soon upon our legs, and although the surface of the snow became worse and worse, and we sank deeper and deeper into it as we proceeded, we managed to do five hours’ work by halting every quarter of an hour. About 3 P.M. I noticed a curious phenomenon. The sun was above the horizon, and was occasionally discernible through the fog—for at this time of the year at this altitude, about 4500 feet, the sun can scarcely be said to set—appearing to move in a circle from the meridian westward, and still keeping above the horizon to almost due north, where it dips for about half-an-hour, appearing again about N.N.E., and by six P.M. it bears due east, some forty degrees above the horizon. A strong current of air was drifting the clouds and fog at our level across the surface of the Jökull from the S.E., while dark masses of cloud were perfectly discernible passing at a very rapid rate across the face of the sun from a precisely opposite direction. The storm now increased in violence, and we were soon so surrounded by whirling clouds of snow that it was impossible to distinguish from what quarter the wind was blowing. The compass had for a long time been almost useless, in all probability owing to the magnetic ore contained in the rocks which underlie the snows of the Jökull. This rendered us entirely dependent upon the wind and the sun for our direction. In clear weather, where the compass is useless, I always steer by a circular piece of card marked off into four right angles, so that by carefully taking the angular bearings of all distinguishable objects, one is able to steer a pretty straight course.
- 42. London, Longmans &Co. E. Weller, Litho. Map of the A u t h o r s R o u t e s from Núpstað to Reikjahlíð
- 43. Being now unableto avail myself of either compass, sun, wind, or card, nothing remained for us but another halt. For two days the storm continued and it would have been impossible to get many yards away from the tent without being lost. On the third day at noon the storm abated, the wind shifted due east, and the sun broke through the clouds. We all turned out, but it was useless to think of struggling through the loose, deep snow. We took our bed out to dry it, for it was wet with the exhalations from our bodies. This, however, was rather against the wish of some of my men, upon whom the inactivity of the last few days had begun to tell. I observed two black peaks protruding through the snow, one about five miles due north, and the other about eleven miles N.W. I was surprised to find a considerable quantity of volcanic ash upon the snow, of a fine, light, grey description. This appeared the more remarkable, as I knew of no volcano that had been in eruption south of the Vatna Jökull, and the storm had blown almost entirely from the S.E. Moreover, I was aware there was no ash of that kind anywhere upon the south. It appeared to me that this must have been carried either from an erupting volcano, or from some ash- strewn district to the north of the Jökull, by a current of air travelling in a different direction to the S.S.E. wind which we had experienced during the last few days, and bisecting the latter current at a point south of our present position, had been unable to resist its force, and had been carried by it to the place where it was now lying. We here obtained an excellent view of the Vatna Jökull Housie, which appeared to be higher than any other point on the Jökull, our present height being 4500 feet—the summit of the Housie being at least 1500 feet above us. Its form is a lop-sided cone, from which I could trace, through my telescope, the course of huge lava streams, now deeply buried in the snow, but still leaving unequal ridges upon each side of the mountain, and in some instances extending to a considerable distance upon the main body of the Vatna Jökull. An extensive eruption of one of these snow-covered volcanoes must be awful, when any vast volume of lava is suddenly ejected upon such a tremendous accumulation of frozen material; but minor eruptions
- 44. and smaller streamsof lava, I should think, can make but little impression upon such an enormous quantity of snow in the first instance. Probably (unless there has been any great amount of sand or ashes previously ejected) they melt their way through the snow to the rocky bed of the mountain, and forming a sort of tube by the aid of the rapidly consolidated crust upon their surface continue their course, much as a lava stream would upon ordinary ground, or more especially, perhaps, at the bottom of the sea, without occasioning any very remarkable phenomena, and even the effect of the most extensive eruptions must of necessity be but local. By 4 P.M. the wind shifted back to its old quarter, S.S.E., and, despairing of frost, we again betook ourselves to the tent. Towards midnight, for about the twentieth time, I went out with Paul to look at the weather. We tried the sleighs, and found it was as much as one man could do to pull a sleigh with nothing on it, and a very small weight almost buried the sleigh in the snow, and enabled it to resist our united efforts to get it along. During our experiment we sunk very deep into the snow. For the last three days I had put every one on half rations, and as anything is better than inactivity with insufficient food, we determined to abandon our sleighs and attempt to force our way through the snow, carrying everything upon our backs. It was rather foggy and sleeting, but the wind was blowing pretty steadily. We communicated our determination to the rest of our party, and they quietly accepted it without a murmur. We packed up everything, and leaving our sleighs and a gathering storm behind us, we turned our faces northward with a cheer which was more animated than might have been expected under the circumstances. I must say our position bore rather a forlorn aspect. Six men heavily laden, wading through snow up to their knees at every step, no view but an ever-advancing circle of gloom, the only variation being that it was darker towards the south, from which quarter a strong wind was blowing, with squalls of sleet and snow. About every quarter of an hour we had to stop from sheer exhaustion, and after two or three hours’ arduous toil two of my men became quite incapacitated and too ill to proceed. This was
- 45. evidently not acase of sham. I therefore halted, and served out with all speed some warm grog; one man was spitting blood, and another was suffering severe pains in the stomach. I had previously advised every man to wear a cloth bandage round his stomach, but none of them had cared to do so. I suffered rather from pains in the bowels the previous year upon the Vatna Jökull, but I was now wearing an abdominal bandage of tarred cloth, and throughout our prolonged stay upon the snow suffered no inconvenience whatever. The next day was finer, with sunshine and increased cold, with snow at intervals, the thermometer being below freezing point all day; one of the sick men had recovered, but the other was still too ill to travel. Towards evening the wind blew from the west, and it began to freeze hard. I therefore sent back for the sleighs, which we had taken the precaution to stand upright and fix firmly in the snow before we left them. By 9 P.M. it was freezing very sharply. I served out an allowance of warm grog, and as the invalid was greatly recovered and said he would rather die than go back, we again struck N.N.E., allowing him to go free. We had packed everything on one of the sleighs, four pulling and one pushing behind, and so firm a crust had now formed upon the surface of the snow that this heavily laden sleigh travelled as easily as an empty one would have done the evening before. We now gradually ascended until at 1 A.M. we reached a rolling plain, at the height of 5750 feet. It was perfectly clear in the west, and I obtained a good view of Tungufell’s and Arnarfell’s Jökulls, which from the angle they made with our line of march, showed me we were two-thirds of the way across the Vatna Jökull. It was still very thick in the south and east, and the wind had shifted to the south- west. An ice-storm was almost the immediate result, a driving mist encrusting everything with ice; the undulations in the plateau became more and more marked, the variation in altitude being sometimes as much as 100 feet or more. A most obdurate mist continued to prevent our obtaining any further view, which was very exasperating, as we might have passed within a short distance of objects of interest without being conscious of the fact.
- 46. We made ourfirst halt at 3 A.M., and took a light meal of Peek and Frean’s meat biscuits and snow. When I say snow I do not mean the pure white frosty snow which lies upon the surface, but the coarse, granular, icy particles of which the crust we were walking upon was composed. I have often been dependent upon snow for the water supply, both in North-West America and upon mountains, and I find the coarser the snow is, and the more it approaches the character of ice, the better it quenches the thirst, and the less likely it is to occasion pain in the stomach. When the fine white snow only can be procured, as every tyro knows, it can be made more palatable by compressing it into a snowball. In other words, the less cold air is swallowed, entangled in the snow, the better; for the very act of squeezing the snow causes it to part with some portion of air, as is shown by the change of colour, as it regelates towards the form of ice. Thus we preferred the coarse icy granules, which formed the crust upon the older snow, to the pure white tempting frost-snow which, owing to the extra amount of air it contained, must have been of a considerably lower temperature than the granulated snow beneath. We were now at the height of 5900 feet, and the temperature was 15° of frost. The rim of the sun was occasionally observable through the fog which surrounded us, giving us a good line to steer by, and bright fog-bows escorted us to windward; but these were simply bows, and had none of the cruciform corona in the interior, which were so observable upon the Myrdal’s Jökull last year. At 6 P.M. we reached a steep ascent, where our compass twisted and turned about in the most eccentric fashion; the heavens became black as night to windward, the wind had risen, and was making the peculiar booming noise I have often remarked in these regions before a storm, and driving a blinding, pitiless drifting snow before it, which eddied about the sleigh and wrapped itself around us, as if longing to enshroud and bury us in its frozen toils. But we had an idea of burying ourselves in our own fashion. “Oskôp mikill stormur kèmur bráðum” (A bad storm is coming on presently), said Eyólfur, sitting down for a moment on the sleigh, and clapping his feet together to knock off the snow which was clinging to his legs, and we were all of the same opinion. We were at the height of 6150
- 47. feet, so Iordered a hole to be dug, and the tent to be pitched. The snow was very hard and firm, even at the depth of four feet, and we cut out as clean a hole as if it had been in salt, but the wind drifted so much loose snow into it, that the men were obliged to hold up the tent to windward during its completion. We had barely got ourselves snug and commenced breakfast, when the storm burst upon us, seeming to threaten the tearing up of the very snow in which we had taken refuge; and had not former experience taught us to fortify our tent well all round with banks of snow, I have no doubt it would have been the last we should have seen of that article of furniture. Being satisfied that all was snug, and that the worst which could happen to us was that we might be buried a few feet in the snow, we went to sleep. When we awoke at mid-day the storm had subsided and the fog had lifted, showing three dark mountains to the north—doubtless Skjaldbreið, Herðubreið, and Dyngjufjöll. We were speculating as to whether we should go on in spite of the still threatening aspect of the weather, when the fog returned, and the booming wind announced another storm to be close at hand. Presently it broke upon us; never before had I heard the wind make such an unearthly wail. It seemed as if every imaginable demon and all the storm spirits of that wild region had assembled to howl and make a united attack upon us. The light was fast becoming obscure, and we were getting fairly snowed up, but that made us all the warmer, all the more secure, and the shrieking of the storm was deadened by the friendly covering. We partook of some chocolate, smoked and sung, and finally slept again. At 8 P.M. the storm had somewhat subsided, and I sent out a man to clear away some of the snow from the roof of the tent to let a little light in. The snow had drifted nearly over the tent, and it took some hard work before we were dug sufficiently out to let in enough light to write by; outside there were 10° of frost, but we were comfortably warm in the tent. The air outside was so full of snow that we could not see a couple of yards in advance. Another day showed us only a continuation of storm and snow which utterly prevented progress. We had now only
- 48. about a week’sprovision left, so I again put every one on half rations. The men were obliged to take turns in clearing away the snow, at intervals of every three hours, from the top of the tent, and before very long the tent had the appearance of lying at the bottom of a deep hole in the snow. We passed the time as best we could, by sleeping, eating, smoking, writing, singing, spinning yarns, and I occasionally amused the assembly by learning strings of Icelandic words by Mr. Stokes’s method of mnemonics, and repeating them in order, either backwards or forwards, which puzzled the Icelanders not a little. Before I started for the Vatna in 1871, I remember saying I should like to see one of its worst storms: I now had that gratification. Storms are interesting natural phenomena, but when prolonged indefinitely are, to say the least, tedious hindrances to progress; and now, lying upon the top of the Vatna Jökull, with the possibility of their lasting for a month, and provisions materially diminishing, their dreary monotony became intolerably oppressive, and after mature consultation we all came to the conclusion that if the weather did not clear in two days’ time, we would leave all impedimenta behind, except provisions, instruments and my diary, and strike northward, storm or no storm—“sauve qui peut.” When we lay down and were fairly snowed over, the booming of the storm sounded as if it came from the interior of the mountain, and almost any familiar sound could be singled out from the hurly- burly in an exaggerated degree, without any great stretch of imagination. It stormed all night; the wind “Trolls” shrieked around us, the thunder of the storm roared through the, to us, dark midnight hours, surging upon the icy bosom of the Jökull, sweeping up its snowy slopes, bearing with it avalanches of snow-drift which had buried us several feet deep by morning. By 5 A.M. it lessened somewhat, the furies of the Vatna appeared to have given up the idea of overwhelming us, and the disheartened tempest sunk away in melancholy sobs, but a determined drift and south-west wind persevered in harassing us.
- 49. It was clearthat we must now start forward, for not only was there a considerable amount of snow yet to be traversed, but a howling wilderness of volcanic sand, lava, and mountain torrents had to be crossed which lay between the north base of the Jökull and the nearest habitation. We could not remain in our present position, so deeply were we buried, and so difficult was it to get in and out of the tent; moreover the fury of the storm had beaten the snow hard, so there was no time to be lost. I served out a hearty meal, and as packing up under such circumstances seemed to demand some stimulant, I made some grog out of methylated spirit, for all our whisky was gone. This served to quicken our circulation, although it was far from being palatable, having, as my Icelanders said, “slœmr dropi,” or a bad after-taste, and no wonder, as the first taste was not suggestive of an agreeable sequel. We packed, but with great difficulty, owing to everything being frozen quite hard. Upon leaving, I drew over my mocassins a pair of fishing stockings; they were as hard as sheet iron, and were a very great inconvenience to me; but it was too cold to stop and take them off, for it seemed as if we should freeze as we stood. These stockings had been of great service in keeping me dry hitherto, and I hoped they would protect me now. I felt a hard lump in the bottom of my left stocking; if it was snow it meant a frozen foot. But there was no help for it—we could not think of stopping to change foot-gear in such a tempest. The wind had shifted to the west, almost freezing the side exposed to it. We steered N.N.E.: it was fortunate the wind was almost at our back, for we could hardly have faced it. After three hours’ hard tugging we reached the height of 6,150 feet, and straight away began to descend, and presently at so rapid a rate that I had to send three men behind, in order to prevent the sleigh from starting on its own account for the bottom of the mountain. Suddenly the clouds cleared away before us, disclosing a deep, snowy valley at our feet, and a tall black mountain, streaked with snow, upon our left and west. Lower and lower we descended, more and more precipitous, till it was evident that we could go no farther upon our present course with the sleigh; so Paul and I went
- 50. forward to explore.The side of the valley terminated in almost perpendicular walls of snow, which were now frozen perfectly hard, and glazed over by the severity of the frost; the opposite side was more broken, with dark crags here and there protruding, while a copious lava stream appeared to flow northwards from the termination of the snow, though I afterwards found that a fringe of glaciers intervened. We next decided on striking due north, along the sloping sides of the valley, to what we supposed to be Querkfjall, but afterwards found to be Kistufell. Upon returning to the sleigh, while putting back my field-glass, which I was obliged to do barehanded, for my gloves were a mass of ice upon the outside, my fingers began to freeze; but a little hard clapping, and by getting two of my men to beat them with their hands, the circulation was restored. I now ordered three of my men to put spiked iron clamps upon their feet, for without this precaution I doubt not but we should have ended our career, sleigh and all, by an abrupt descent into the valley beneath, unless we had been stopped by some of the ugly crevasses which yawned half-way down the snowy steep, upon the slippery and precipitous sides of which we were descending. We proceeded, but with great difficulty; our trouble now being, not that the sleigh was hard to get along, but that it would go too fast; in fact, it seemed likely to run away with us altogether. Behind us was a fierce wind, beneath us a precipice of some 800 or 1,000 feet; and the sloping snow-banks we were treading shelved off at such an alarming angle that it rendered the work more dangerous than pleasant. In this critical position I became painfully aware that I had frozen my left big toe; for the increased exertions and the lessening altitude were causing it to thaw. The pain was horrible; but presently the slope became less abrupt, and we stepped along at such a rate that 1,500 feet were negociated with considerable speed. Hurrah! we were again in bright sunshine; but the moment we stood still, the wind cut us to the bone. Before us lay the long looked-for Norðurland. We arrived at the bottom of the valley, and found it full of loose snow, which was knee-deep, for the crust was
- 51. here much toolight to bear our weight, and at every few feet we sunk into a miniature crevasse. After struggling on for some few hours, however, we pitched our tent. Right thankful was I to get some warm soup and creep into the bag. One of my men—and a real good fellow he was—named Sigurð, cuddled my left foot in his arms, although my writhings kept him as well as myself awake while the others slept. I have had many parts of my body frozen, but I never suffered so much as from that toe. After a few hours we again started; and although the sleigh travelled easily over the crust, we still broke through it, which occasioned me so much pain at every step that I sat upon the sleigh and was drawn along until we had descended so much that the crust ceased altogether. The snow terminated in a half-melted slush, lying upon a bottom of ice. Wading through the slush, which at times took us up to the waist, we next reached Kistufell, where the ice and snow terminated. Here we landed on a bed of volcanic débris, which covered the ice to such a depth that one could in no way, except by digging, distinguish it from the adjacent fjall. The Vatna Jökull now lay behind us with its mysterious recesses and volcanoes carefully guarded from intrusion by gloom and storm. To the north of us rose a cluster of mountains from which great quantities of steam were rising, and hovering above their summits in a huge mushroom- shaped cloud; to our left and west lay a wide-spreading lava-field, arms of which stretched amongst the neighbouring mountains like the troubled waters of a cindery ocean. Patches of black sand at intervals broke the continuity of this tract of lava, and culminated in a desert still farther to the north-east. Beyond, all the weird forms of fire-wrought mountains formed a fitting back-ground; their rude outlines rendered still more uncouth and grim by the fierce storms of ages. A huge tongue of glaciers at this point swept down to a distance of some ten miles beyond its most northern limit, as represented upon the map published by Olsen in 1844, from a survey made by Gunnlaugsson, in 1835. I here caught sight of Snæfell; and, upon taking its bearings with the smoking mountains,
- 52. which were evidentlythe Dyngjufjöll, I found that instead of being at Querkfjall, which was the point I had intended to strike, we were upon the east side of Kistufell, about ten miles too much to the west. What astonished us most was (granted that we were at the east side of Kistufell) that we could see nothing of the Jökulsá-á- fjöllum, which river, upon Olsen’s map, rises at the foot of Kistufell; besides, upon his map the Jökull ends at Kistufell, while here a huge glacier extended east and north-east as far as the eye could reach, though exactly to our north and north-west it terminated abruptly, and only an insignificant river flowed to the north. We here abandoned our sleigh and snow-shoes which had served us so well, and whatever we no longer required, and, making everything into packs, continued our descent over huge piles of moraine, which doubtless covered glacial ice, buttresses and points of which here and there protruded. Having slidden down several steep slopes of snow, which had collected in all the hollows, affording us ready means of descent, we found ourselves at the height of 3,850 feet, in the bed of what evidently had been a large river, though now only an insignificant stream. To our east and right stretched the immense glaciers before mentioned, completely overrunning the route taken by Gunnlaugsson in 1835, and diverting the source of the Jökulsá, which rises in several arms from the extremity of the glacial tongue before mentioned. Upon our left and west lay the wide-spreading lava-desert of the Ódáðahraun. Our way over the sandy bottom of the grand old watercourse was an easy one to travel, for the sand had absorbed sufficient water to make it firm and compact. Our attention was engaged for some time in watching the fanciful shapes that crowned the dark wall of ice upon our right, on the opposite side of the stream which now lay between us and the glacier; and now and then we could not help stopping to peer into some of the dark chasms which seemed to penetrate into the heart of the icy monster, and to admire the little cataracts of foam which spouted from clefts in the dark green ice, or to wonder at some icy pinnacle or turret, that ever and again
- 53. tumbled from, perhaps,some few hundred feet above us with a roar and a splash into the river, there to be slowly melted, while the sound of its downfall echoed and re-echoed amongst the cavernous openings in the glacier from which it had fallen! After an hour or so we settled on a low sandy island in the middle of the river, which must have formed formidable rapids when the immense stream that had hollowed out this mighty watercourse had roared over its bed; but it was shallow enough now, and by judiciously picking our way it scarcely reached up to our knees as we waded to the little island. I here noticed, as I had often done before, an intermittent occurrence of waves in certain portions of the stream. These, in large rivers, are rather terrible things, but here they were on so small a scale as to make their examination simply a safe indulgence of harmless curiosity. These waves occur in all the sandy rivers, and they are occasioned by the sand and detritus, which is brought down by the river in large quantities, accumulating against some obstacle until such a time as it forms rapids, which increase in proportion to the durability of this suddenly-formed sandbank. In most cases it readily yields to the action of the water, and is carried away; if, however, the material which is thus piled up should be of a heavier character than usual, it soon accumulates to such an extent as to resist the action of the water altogether, and cause the current to alter its course. This shows how the rivers of Iceland may be diverted and changed from this cause alone, converting shallows into deep water, and deep water into shallows, indeed altering the position and character of the rivers altogether. As we lay down, the volcanoes in the Dyngjufjöll were smoking away with increased violence. My frost-bitten toe would not allow me to sleep much, so after a doze of two hours we started on our way; we had but two days’ full rations left, and as Grímstaðir was the nearest farm, a series of forced marches was necessary. Before us to the N.E. was a cluster of hills, which stretched from the southern extremity of the Dyngjufjöll in a S.E. direction towards the Jökulsá, upon the east and west sides of which valleys appeared to open northwards. Wishing, however, to get a good view of the
- 54. country before us,as neither of us had been here before, and it was a matter of paramount importance that we should make no mistake as to the direction, I decided to steer for the centre of the hills, and to cross them. For a short distance we skirted the tongue of the Jökull, past a line of moraine which shewed that the glacier had ebbed as well as flowed, then bearing more to the north, after a hard walk of three hours we reached the hills before us. They were composed of the usual confusion of agglomerate, sand and lava, which had issued from it—it was impossible to say where; but they were evidently of a very ancient date, and many of the harder rocks were glaciated, while the softer ones were simply ruinous heaps. After an arduous scramble, we crossed these hills and reached the little desert of black volcanic sand we had seen from the northern edge of the Vatna Jökull. This sand plain lay between the Dyngjufjöll and a chain of mountains upon the opposite side of the Jökulsá-á- fjöllum. It was now raining somewhat heavily, but there was no fog; the burdens of my men were heavy, and I was carrying all I could manage with my bad foot. Under these conditions we were obliged often to rest, which much hindered in our progress. We sighted some low, black, misshapen volcanoes, about half way across the plain, and near these we determined to camp for the night. Two hours brought us to a field of lava which had flowed from and surrounded those eccentric little volcanoes which rose in four ghastly eminences in the centre of the plain, in no part more than 100 feet high. Tired as I was, and greatly inconvenienced by my foot, I could not refrain from examining them. They were situated upon a crack from which the lava had welled up in four mamelonic shapes, which in two instances showed irregular breached craters, nearly filled with sand, which had been drifted thither by the wind. The lava was basaltic, and of a remarkably scoriaceous nature, though in the immediate neighbourhood of the volcanoes no cinders were visible around them, so their eruptions must have been attended with but little of explosive character. The worst feature of our night’s lodging was the absence of water, so I ordered the waterproof coats to be spread out to catch rain for
- 55. our use inthe morning. It was 1.30 A.M. before we all turned in for the night. Sand is warm to camp upon, but it gets into everything, and when one is wet it sticks to clothes, &c., in a most objectionable manner. By six A.M. we were all awake, sufficient water had collected for immediate use, and we were soon all under weigh over the lava, which in most places flowed very evenly, and being of a more compact character than that which was close to the volcanoes we had just left, had allowed little pools of rain water to collect upon the surface. We marched for four hours, and then struck a large river upon our east. This was the Jökulsá-á-fjöllum. After following its course for some time, we decided to “cache” everything but the remainder of our provisions, our maps, and my diary, for it was my intention to return with horses to the Dyngjufjöll mountains which now lay to our N.W., when I could recover them without much difficulty. Having carefully made our “cache,” we planted a flag-pole upon an adjacent sandbank, and having carefully taken its bearings, struck for the Vaðalda hills, which were not very far distant. These hills run for some nine miles parallel with the course of the Jökulsá; their base being washed by the Svartá, or Black river, which rises in the Dyngjufjöll, but is soon lost in the sand, re-appearing on the Svartá at the commencement of the Vaðalda. Upon the opposite side of this river we found a root of angelica (Icelandic, hvönn), the stem and root of which we shared and ate with great relish; we also saw two white sheep, but how they manage to eke out an existence must have puzzled their sheeps’ heads not a little. Though, proverbially, two heads are better than one, I doubt if the proverb would hold good in their case, but there may be some grass in glens which I have not seen in the Ódáðahraun, where enough herbage may grow to feed Icelandic sheep, as they are not very dainty, and are accustomed to short commons. The Vaðalda hills, although of no great height, command an extensive view towards the Vatna Jökull, and upon reaching their summit I glanced back over the plain. It was one broad wilderness of black sand and lava, girt about with ridges of volcanic mountains,
- 56. whose numerous conesand chasms have vomited the immense amount of ash, sand and lava with which the surrounding country is covered. In the centre of the plain rose the little volcanoes by which we had encamped the previous night, grimly and perkily protruding, as if they aped their monster brethren around them. Beyond all was the wide, white expanse of the Vatna Jökull, from which a huge tongue of glacier extended more than half way across the plain; from its extremity commenced the river we had been following (the Jökulsá-á-fjöllum), which stretched through the black bare plain sometimes in many arms, enclosing little islands of black sand and pebbles in its sinuous embrace, then surging along through a single deep channel it had worn for itself in the sand, where the unstable banks, even while we gazed on them, were crumbling and falling in, patch after patch of sand rendering still more murky its already discoloured waters. From here I obtained the first good view of the Querkfjall, which appeared to be a cluster of conical mountains, one huge crater being on the northern side of the Vatna Jökull. This large crater, though partially filled with snow, was smoking at three points, but presented no other signs of activity. Having advanced about a mile upon the Vaðalda, we were soon upon the pumice which was ejected last year from the Öskjugjá, or chasm of oval casket, in the Dyngjufjöll mountains. It has fallen in a line of about twenty-five miles in breadth from the centre of the Vaðalda to the south of Herðubreið, in a band of continually extending ladià eastward towards the sea shore, destroying in its course six farms in the Jökuldalr, and injuring others in the immediate vicinity. This shows that the prevalent winds during the eruption of Öskjugjá must have been south-west. This pumice is of a remarkably vitreous nature and vesicular in structure, often assuming very beautiful forms, such as sponge, honeycomb, coral or grained wood. As far as the eye could see, the whole country was buried under greyish cinders, often to the depth of several feet; while in places it had been swept up into huge banks of many feet in thickness by the wind, sometimes burying whole lava fields, the more elevated crags of which protruded, as if struggling
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