The VIII International Symposium on Tardigrada

The congress was attended by 62 researchers from 11 countries from five continents (Africa, Asia, Australia, Europe and North America). During the six days of Symposium 35 talks and 18 posters were presented.

This tardigrade meeting was the only one so far to be held in a Scandinavian country. It was also the last Symposium in the XX century.

Two days after the conference a workshop on Arctic Tardigrades was held at a Danish Arctic Station, Qeqertarsuaq, Greenland from the 7 to 18 August 2000. There were 15 participants taking part in the Greenlandic Adventure.

Table of contents

Group photo


The Symposium logo, designed by Professor Reinhardt M. Kristensen and created by Brigitte Rubaek (University of Copenhagen, Denmark), shows four individuals of a marine tardigrade, Tanarctus bubulubus.


30 July 2000 - 05 August 2000.


Copenhagen, Denmark.

World Map from Wikipedia.


Zoological Museum, University of Copenhagen.


Planning group:
Professor Reinhardt Møbjerg Kristensen (University of Copenhagen),
Dr. Jette Eibye-Jacobsen (University of Copenhagen),
Dr. Hans Ramløv (Roskilde University),
Dr. Aslak Jørgensen (University of Copenhagen),
Anne Marie Richardy Warfield (University of Copenhagen).

Secretarial services:
Anne Marie Richardy Warfield (University of Copenhagen),
Vibeke Boye (University of Copenhagen).

Student volunteers:
Lars Haumann (University of Copenhagen),
Iben Heiner (University of Copenhagen),
Jesper Kragh (University of Copenhagen),
Katrine Worsaae (University of Copenhagen).

Participants (alphabetically)


 Altiero, Tiziana (Italy)
 Bateman, L.
Clark W. (USA)
Bergström, Jan W. (Sweden)
Roberto (Italy)
T.M. (Denmark)
Graham (Sweden)
M. (Germany)
Peter (Australia)
Sandra (Australia)
 Collins, Michael (Canada)
A. (Italy)
D’Addabbo, Maria Gallo (Italy)
William (USA)
Ruth A. (USA)
Karen (USA)
Petra (Germany)
Jette (Denmark)
E. (Denmark)
Silvana (Italy)
James R. (USA)
Margaret (UK)
Philip M. (UK)
Hartmut (Germany)
A. (Denmark)
Roberto (Italy)
Jesper Guldberg (Denmark)
Iben (Denmark)
Melisa (USA)
Høeg, Jens T. (Denmark)
Amber (USA)
Svanhild (Germany)
Masamichi (Japan)
K. Ingemar (Sweden)
Aslak (Denmark)
Ask M. (Denmark)
S.M. (Denmark)
Reinhardt M. (Denmark)
Katrin-Theres (Germany)
Niklas (Denmark)
López, Noemi Guil (Spain)
Andreas (Germany)
Nigel (UK)
Sandra J. (UK)
Harry A. (USA)
Roger (Botswana) 
William Randy (USA)
Nadja (Denmark)
Diane R. (USA)
Jack (USA)
Kenneth J. (USA)
Nielsen, Claus (Denmark)
Thorsten (Denmark)
Lorena (Italy)
 Romano III,
Frank A. (USA)
Hilke (Germany)
Andreas (Germany)
Rolf (Germany)
 Sørensen, Martin Vinther (Denmark)
Anne Marie Richardy (Denmark)
Peter (Denmark)
Jonathan (USA)


 Beasley, Clark W. (USA)
 Bertolani, Roberto (Italy)
Eibye-jacobsen, Jette (Denmark)
Silvana (Italy)
Roberto (Italy)
Hansen, Jesper Guldberg (Denmark)
 Hohl, Amber (USA)
Jönsson, K. Ingemar (Sweden)
Jørgensen, Aslak (Denmark)
Kristensen, Reinhardt M. (Denmark)
Marley, Nigel (UK)
Mcinnes, Sandra J. (UK)
Miller, William Randy (USA)
Nelson, Diane R. (USA)
Warfield, Anne Marie Richardy (Denmark)

Group photo

Photo by Geert Brovad.



Sunday (30.07.2000) 
August Krogh Institute - Universitetsparken 13
19:00-20:00 Start mounting posters
20:00-22:00 Video from the Ice Cap of Greenland and Icebreaker Party


Monday (31.07.2000) 
August Krogh Institute - Universitetsparken 13
09:00-10:30 Registration at August Krogh Institute, Universitetsparken 13
10:30-10:45 Dean of the Science Faculty, H. Jeppesen
10:45-11:00 Director of the Zoological Museum, H. Enghoff
11:00-11:30 Morning tea or coffee
Chairman: D.R. Nelson
11:30-12:30 Garey, J. & Nichols, P.B.  Keynote lecture:
Molecular analysis and tardigrade systematics.
12:30-14:00 Lunch
14:00-14:30 Nielsen, C.  Panarthropods: articulates or ecdysozoans?
14:30-15:00 Sørensen, M.V. Modern computer cladistics: Cladogram of the Animal Kingdom.
15:00-15:30 Afternoon tea or coffee
Chairman: H. Greven
15:30-16.00 Schmidt-Rhaesa, A. Tardigrades - are they really miniaturized dwarfs?
16:00-16:30 Dewel, R.A. & Dewel, W.C. A new hypothesis for the origin of the Bilateria: Implications for understanding the body plan of tardigrades.
16:30 Bus departure for Town Hall
17:00-18:00 Reception at the Copenhagen Town Hall


Tuesday (01.08.2000) 
August Krogh Institute - Universitetsparken 13
Chairman: C.W. Beasley
10:00-10:30 Romano, F.A.III, Barreras-Borrero, B. & Nelson, D.R.  Ecological distribution and community analysis of Tardigrada from Choccolocco Creek, Alabama, with the description of a new species of Echiniscus.
10:30-11:00 Claxton, S.K. Antechiniscus in Australia: Description of Antechinicus sp.n. and redescription of Antechiniscus parvisentus (Horning & Schuster, 1983) (Heterotardigrada: Echiniscidae).
11:00-11:30 Morning tea or coffee
11:30-12:00 Nichols, P.B., Romano, F.A.III, & Nelson, D.R.  Seasonal and altitudinal variation
in the distribution and abundance of Tardigrada on Dugger Mountain, Alabama.
12:00-13:00 Round Table Discussion: Techniques for whole-mount preparations of Tardigrada.
Chairs: Bertolani, R., Claxton, S.K., Galio D'Addabbo, M., and Mclnnes, S.J.
13:00-14:00 Lunch
Chairman: J. Eibye-Jacobsen
14:00-14:30 Bertolani, R.  Evolution of reproductive mechanisms in tardigrades.
14:30-15:00 Greven, H. & Kristensen, R.M. Notes on spermiogenesis of
Echiniscoides sigismundi.
15:00-15:30 Altiero, T., Rebecchi, L. & Bertolani, R. Banding techniques on tardigrade chromosomes.
15:30-16:00 Afternoon tea or coffee
Chairman: R.A. Dewel
16:00-16:30 Rebecchi, L.  Spermatozoon in tardigrades: Evolution and relationships with
the environment.
16:30-17:00 Guidetti, R. & Bertolani, R. New data on the phylogeny of the Macrobiotidae (Tardigrada, Eutardigrada).
Greaves, P.M. Royal tardigrades - The fauna of Buckingham Palace, London.
Greven, H. & Schüttler, L. How to crawl and dehydrate on moss.
Guidetti, R. & Bertolani, R. Long-term ecological studies on tardigrades in a beech leaf Litter.
Guidetti, R. & Bertolani, R. Tardigrades of Piane di Mocogno (Northern Apennines, Italy).
Ito, M. & Abe, W. Depth distribution of soil inhabiting tardigrades in a subalpine coniferous forest of Japan.
Meyer, H.A. Tardigrades of Arkansas and Louisiana, U.S.A.
Miller, W.R. & Case, S.B. KanCRN: a collaborative research model using tardigrades.
Miller, W.R. & Miller, J.D. Return to Terra Incognita: more giant tardigrades.
Peters, T. & Dumjahn, P. Moss-living tardigrades from two altitudinal transects at Disko lsland, West Greenland.


Wednesday (02.08.2000) 
August Krogh Institute - Universitetsparken 13
Chairman: W.C. Dewel
09:00-10:00 Ruhberg, H.  Keynote lecture:
Recent Onychophora: are they a sister group of the Tardigrada?
10:00-10:30 Kristensen, R.M., Møbjerg, N. & Jørgensen, A. The cephalic sensory structures
and the brain of Actinarctus doryphorus (Arthrotardigrada).
10:30-11:00 Eibye-Jacobsen, J. A new method for making SEM-preparations of the tardigrade buccopharyngeal apparatus.
11:00-11:30 Morning tea or coffee
Chairman: W.R. Miller
11:30-12:00 Bateman, L. & Collins, M. Tardigrades of Newfoundland, Canada.
12:00-12:30 Collins, M. & Bateman, L. The biogeography of tardigrades of Newfoundland, Canada.
12:30-13:00 Nelson, D.R. & Adkins, R.G. Distribution of tardigrades within a Moss Cushion: Do tardigrades migrate in response to changing moisture conditions?
13:00-14:00 Lunch
14:00-14:30 Marley, N.J. Tardigrade assemblages from an altitudinal transect in Venezuela.
14:30-15:00 Nickell. K.J. & Miller, W. R. Tardigrades of South America: Macchu Picchu and Ollantaytambo, Peru.
15:00-15:30 Afternoon tea or coffee
Altiero, T. & Rebecchi, L. Results and problems in rearing tardigrades.
Brockmann, C. & Ruhberq. H. Ooperipatellus decarafus (Peripatopsidae):
Hatching observations in an oviparous onychophoran.
De Zio Grimaldi, S. & Galla D'Addabbo, M. Further data on the Mediterranean Sea tardigrade fauna.
Jönsson, K.I., Borsari, S., & Rebecchi, L. Do populations within tardigrade species differ in anhydrobiotic capacity?
Jørgensen, A. A review and graphical presentation of the African tardigrade fauna using GIS with the description of Isohypsibius nov.sp. (Eutardigrada: Hypsibiidae) from Lake Malawi.
Mclnnes, S.J., Chown, S.L., Dartnall, H.J.G., & Pugh, P.J.A. Milnesium cfr tardigradum a monitor of high altitude micro-invertebrates on sub-Antarctic Marian Island.
McInnes, S.J. Is it real?
McInnes, S.J. Exposed: Danger lurking in the Maritime Antarctic freshwater algal mats for rotifers and tardigrades.
Middleton, R. Ecology of tardigrades in southern Africa: A preliminary report.
Russell, P.M., Marley, N.J. & Hockings, M.E. Do confocal microscopy and tardigrades have a future together?


Thursday (03.08.2000) 
09:30 Bus departure from August Krogh Institute.
10:00-12:00 Visit to Louisiana Museum of Modern Art.
12:00-12:30 Bus drive to Kronborg (The castle of Hamlet).
12:30-14:00 Lunch at Kronborg Castle.
14:00-16:00 Sightseeing on the site of Shakespeare's Hamlet and the Casemates.
16:00-16:30 Bus drive to Hillerød.
16:30-17:30 Sightseeing: the fountains, gardens and castle of Frederiksborg.
17:30-18:00 Bus drive back to Copenhagen.


Friday (04.08.2000) 
August Krogh Institute - Universitetsparken 13
Chairman: H. Ruhberg
09:00-10:00 Bergström, J. & Hau, X.-g.  Keynote lecture:
Cambrian Onychophora or Tardipolypoda.
10:00-10:30 Budd, G.  Tardigrades as "stem-group" arthropods: The evidence from the Cambrian fauna.
10:30-11:00 Maas, A. & Waloszek, D. Fossil stemline arthropods -  tardigrades, lobopodes and pentastomids from the Cambrian - An 'Orsten' perspective.
11:00-11:20 Morning tea or coffee
Chairman: G. Budd
11:20-11:50 Jørgensen, A. & Kristensen, R.M.  A new tanarctid arthrotardigrade with buoyant bodies.
11:50-12:20 McInnes, S.J. Biodiversity and biogeography of the global marine tardigrade fauna.
12:20-12:50 Boesgaard, T., Jørgensen, A. & Kristensen, R.M. Tardigrades from Australian
marine caves.
12:50-13:10 Hansen, J.G., Jørgensen, A. & Kristensen, R.M. Preliminary studies of the tardigrade fauna of the Faroe Bank.
13:10-14:00 Lunch
Chairman: K.J. Nickell
14:00-14:30 Hohl, A.M. & Miller, W.R.  Tardigrades of North America: As possible bio-indicators relative to a power plant.
14:30-15:00 Herbert, M. & Miller, W.R. Tardigrades of North America: A study of populations in Southern Indiana.
15:00-15:30 Afternoon tea or coffee
15:30-16:00 Symposium photograph
16:00-17:00 Round Table Discussion: The phylogeny of Tardigrada
Chairs: Bertolani, R., Budd, G., Greven, H., Dewel, R.A. & Nelson, D.R.
19:00-21:00 Symposium Dinner at Tivoli Gardens


Saturday (05.08.2000) 
August Krogh Institute - Universitetsparken 13
Chairman: R. Bertolani
09:00-10:00 Wright, J.  Keynote lecture:
Cryptobiosis 300 years on from van Leeuwenhoek: What have we Learned about tardigrades?
10:00-10:30 Jönsson, K.I. & Rebecchi, L. Natural selection on cryptobiotic capacity in tardigrades.
10:30-11:00 Ramløv, H. & Westh, P. Aspects of cryptobiosis in the eutardigrade Adorybiotus
(Richtersius) coronifer.
11:00-11:20 Morning tea or coffee
Chairman: H. Ramløv
11:20-11:50 Westh, P.  Bound water and cryptobiosis.
11:50-12:20 Beasley, C.W. Response of Macrobiotus hufelandi (Tardigrada) to light.
12:20-14:00 Lunch
14:00-15:00 Kristensen, R.M. (convener)



The Symposium talks have been published in a special volume of 'Zoologischer Anzeiger'. The volume contains 41 scientific papers on tardigrade morphology, taxonomy, phylogeny, evolution, biogeography, faunistics, ecology, physiology, reproduction, behaviour and methodology. The symposium proceedings are dedicated to the memory of Professor Mario Bertolani.

Zoologischer Anzeiger
A Journal of Comparative Zoology)
vol. 240(3-4), 2001

Proceedings of the Eighth International Symposium on Tardigrada

Reinhardt Møbjerg Kristensen
(Guest Editor)






(click paper titles to open full texts in PDF)



Kristensen, R.M. Foreword. 213-215
Jeppesen, H. Welcome by the Dean Henrik Jeppesen at the 8th International Symposium on Tardigrada. 216
Altiero, T. & Rebecchi, L. Rearing Tardigrades: Results and Problems. 217-221
Bateman, L. & Collins, M.A. Preliminary Account of the Tardigrades of Newfoundland. 223-232
Beasley, C.W. Photokinesis of Macrobiotus hufelandi (Tardigrada, Eutardigrada). 233-236
Bergstrom, J & Hou, X.G. Cambrian Onychophora or Xenusians. 237-245
Bertolani, R. Evolution of the Reproductive Mechanisms in Tardigrades - A Review. 247-252
Boesgaard, T.M., Kristensen, R.M. Tardigrades from Australian Marine Caves. With a Redescription of Actinarctus neretinus (Arthrotardigrada). 253-264
Budd, G.E. Tardigrades as 'Stem-Group Arthropods': The Evidence from the Cambrian Fauna. 265-279
Claxton, S.K. Antechiniscus in Australia: Description of Antechiniscus moscali sp. n. and Redescription of Antechiniscus parvisentus (Horning & Schuster, 1983) (Heterotardigrada: Echiniscidae). 281-289
Collins, M. & Bateman, L. The Ecological Distribution of Tardigrades in Newfoundland. 291-297
Dastych, H. Notes on the Revision of the Genus Mopsechiniscus (Tardigrada). 299-308
Eibye-Jacobson, J. A New Method for Making SEM Preparations of the Tardigrade Buccopharyngeal Apparatus. 309-319
Garey, J.R. Ecdysozoa: The Relationship between Cycloneuralia and Panarthropoda. 321-330
Greven, H. & Kristensen, R.M. Some Aspects of Spermiogenesis and Spermatozoan Ultrastructure in Echiniscoides sigismundi (Heterotardigrada: Echiniscoididae). 331-339
Greven, H. & Schuttler, L. How to Crawl and Dehydrate on Moss. 341-344
Grimaldi, S.Z. & D'Addabbo, M.G. Further Data on the Mediterranean Sea Tardigrade Fauna. 345-360
D'Addabbo, M.G., Grimaldi, S.Z. & Sandulli, R. Heterotardigrada of Two Submarine Caves in S. Domino Island (Tremiti Islands) in the Mediterranean Sea with the Description of Two New Species of Stygarctidae. 361-369
Guidetti, R. & Bertolani, R. Phylogenetic Relationships in the Macrobiotidae (Tardigrada: Eutardigrada: Parachela). 371-376
Guidetti, R. & Bertolani, R. The Tardigrades of Emilia (Italy). III. Piane di Mocogno (Northern Apennines). 377-383
Hansen, J.G., Jorgensen, A. & Kristensen, R.M. Preliminary Studies of the Tardigrade Fauna of the Faroe Bank. 385-393
Hohl, A.M., Miller, W.R. & Nelson, D.R. The Distribution of Tardigrades Upwind and Downwind of a Missouri Coal-Burning Power Plant. 395-401
Ito, M.T. & Abe, W. Micro-distribution of Soil Inhabiting Tardigrades (Tardigrada) in a Sub-alpine Coniferous Forest of Japan. 403-407
Jonsson, K.I. The Nature of Selection on Anhydrobiotic Capacity in Tardigrades. 409-417
Jonsson, K.I., Borsari, S. & Rebecchi, L. Anhydrobiotic Survival in Populations of the Tardigrades Richtersius coronifer and Ramazzottius oberhaeuseri from Italy and Sweden. 419-423
Jorgensen, A. & Kristensen, R.M. A New Tanarctid Arthrotardigrade with Buoyant Bodies. 425-439
Jorgensen, A. Graphical Presentation of the African Tardigrade Fauna Using GIS with the Description of Isohypsibius malawiensis sp. n. (Eutardigrada: Hypsibiidae) from Lake Malawi. 441-449
Maas, A. & Waloszek, D. Cambrian Derivatives of the Early Arthropod Stem Lineage, Pentastomids, Tardigrades and Lobopodians An 'Orsten' Perspective. 451-459
McInnes, S.J., Chown, S.L., Dartnall, H.J.G. & Pugh, P.J.A. Milnesium cfr. tardigradum (Milnesiidae, Apochela, Tardigrada): A Monitor of High Altitude Meiofauna on Sub-Antarctic Marion Island. 461-465
McInnes, S.J. Is It Real? 467-469
Meyer, H.A. Tardigrades of Louisiana and Arkansas, United States of America. 471-474
Miller, W.R., Horning, D.S., Heatwole, H. F. Tardigrades of the Australian Antarctic: Macquarie Island, sub-Antarctica. 475-491
Nelson, D.R. & Adkins, R.G. Distribution of Tardigrades within a Moss Cushion: Do Tardigrades Migrate in Response to Changing Moisture Conditions? 493-500
Nichols, P.B., Romano III, F.A. & Nelson, D.R. Seasonal and Altitudinal Variation in the Distribution and Abundance of Tardigrada on Dugger Mountain, Alabama. 501-504
Nickel, K., Miller, W.R. & Marley, N.J. Tardigrades of South America: Machu Picchu and Ollantaytambo, Peru. 505-509
Pilato, G. & Binda, M.G. Biogeography and Limno-terrestrial Tardigrades: Are They Truly Incompatible Binomials? 511-516
Ramlov, H. & Westh, P. Cryptobiosis in the Eutardigrade Adorybiotus (Richtersius) coronifer: Tolerance to Alcohols, Temperature and de novo Protein Synthesis. 517-523
Rebecchi, L. The Spermatozoon in Tardigrades: Evolution and Relationships with the Environment. 525-533
Romano III, F.A., Barreras-Borrero, B. & Nelson, D.R. Ecological Distribution and Community Analysis of Tardigrada from Choccolocco Creek, Alabama. 535-541
Russell, P.M., Marley, N.J. & Hockings, M.E. Do Confocal Microscopy and Tardigrades Have a Future Together? 543-548
Schmidt-Rhaesa, A. Tardigrades - Are They Really Miniaturized Dwarfs? 549-555
Liltorp, K., Jakobsen, K.L., Nielsen, O.F., Ramlov, H. & Westh, P. Bound Water and Cryptobiosis: Thermodynamic Properties of Water at Biopolymer Surfaces. 557-562
Wright, J.C. Cryptobiosis 300 Years on from van Leuwenhoek: What Have We Learned about Tardigrades? 563-582



Click here to download the booklet with the programme, abstracts and other Symposium info.


By Reinhardt M. Kristensen.

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