9th International Symposium on Tardigrada St. Pete Beach, FL 28 July – 1 August 2003
Program of Events
Date/Time Event Location______
Sunday July 27, 2003
6:00pm – 8:00pm Welcome Social Beachfront Breezeway
Monday July 28, 2003
Session 1 Welcome Mediterranean Palm
9:00am Jim Garey, Symposium Committee Chair
9:10am Sandy Schneider, Associate Dean of Research, USF
9:30am Diane Nelson, East Tennessee State University Tardigrade Research: Where Have We Been? Where Are We Going?
10:30am Morning Break Mediterranean Palm
Session 2 Phylogeny Mediterranean Palm
11:00am Jerome Reiger, University of Maryland
12:00 – 1:30pm Lunch Royal Palm – 8th Floor
1:30pm Mark Blaxter, University of Edinburgh Molecular phylogenetics of the Tardigrada and an investigation of the position of tardigrades in animal phylogeny
2:00pm Jette Eibye-Jacobsen, University of Copenhagen Three dimensional understanding of the foremost section of the buccal apparatus of Echiniscus viridissimus Peterfi, 1956 (Heterotardigrada)
2:30pm Ruth Dewel, Appalachian State University Origin and Diversification of the Arthropods: New Interpretations of Some Old Characters
3:00pm Afternoon Break Mediterranean Palm
3:30pm Roberto Bertolani, University of Modena Phylogenetic relationships in Macrobiotidae (Tardigrada). II. Molecular (mtDNA) and morphological approaches
4:00pm P. Brent Nichols, University of South Florida Family Values: A Cladistic Analysis of the Tardigrada 9th International Symposium on Tardigrada 2
Date/Time Event Location______
Tuesday July 29, 2003
Session 1 Life History – Temperate Mediterranean Palm
9:00am Tiziana Altiero, University of Modena Phenotypic life history variations in two clones of Macrobiotus richtersi
9:30am Harry Meyer, McNeese State University Distribution of Terrestrial Tardigrades in the State of Florida
10:00am Nigel Marley, University of Plymouth Preliminary Results from a Study on Ecuadorian Tardigrada
10:30am Morning Break Mediterranean Palm
11:00am Juliana Hinton, McNeese State University Seasonal and Spatial Variation in Tardigrade Diversity in Leaf Litter from Florida and Louisiana
11:30am Paul Bartels, Warren Wilson College A Large-scale, Multi-Habitat Inventory of Tardigrades in the Great Smoky Mountains National Park.
12:00 – 1:30pm Lunch Royal Palm – 8th Floor
Session 2 Poster Session 1 Sabal - Canary Palm 3rd Floor
1:30pm Wataru Abe, Hokkaido University Semiterrestrial Tardigrades form Sakhalin Island, Far East Russia
Jennifer Daub, University of Edinburgh Genomic Resources for the Tardigrade Hypsibius dujardini
Peter Degma, Comenius University The Ecological Distribution of Tardigrada in National Nature Reserve Stužica (Bukovské vrchy Mts , NE Slovakia )
Maria Fernandez, Universidad Nacional de La Pampa Population Dynamics of Dactylobiotus grandipes Schuster et al., 1977 (Tardigrada) in a Neotropical Eutophic Pond.
Roberto Guidetti, University of Modena Dactylobiotus octavi n. sp. (Eutardigrada; Macrobiotidae) from Disko Island
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 3
Date/Time Event Location______
Session 2 Poster Session 1- cont. Sabal - Canary Palm 3rd Floor
Jesper Hansen, University of Copenhagen The “Hyena Female” within the Marine Tardigrada with the Description of Two New Species of Megastygarctides (Arthrotardigrada: Stygarctidae) from Saudi Arabia
Nigel Marley, University of Plymouth Designation of Pseudobiotus kathmanae Nelson (Tardigrada) as the Type Species of Pseudobiotus Nelson.
Daiki Horikawa, Hokkaido University The Effects of Prehydration on the Anhydrobiotic Survival in the Tardigrade Milnesium tardigradum
Javier Jerez-Jaimes, University of Puerto Rico Tardigrades in Six Phorphytes of the Moss Calymperes tenerum C. Müller
ElianaNarvaea, University of PuertoRico Tardigrade Community Composition in Four ForestTypes in the El Divisio Reserve, (Santander, Colombia)
Atsushi Suzuki, University of Keio Oogenesis of Milnesium tardigradum
Sandra McInness, British Antarctic Survey Exceptional, Tardigrade Dominated Ecosystems in Ellsworth Land, Antarctica.
Diana Sucharski, Chestnut Hill College Tardigrades of North America: Southeastern Pennsylvania
3:00pm Afternoon Break Mediterranean Palm
3:30pm FreeTime/Late Additions
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 4
Date/Time Event Location______
Wednesday July 30, 2003
Session 1 Life History – Polar and Sub-polar Mediterranean Palm
9:30am Michael Collins, University of Newfoundland A Preliminary Account of Tardigrades of Labrador, Canada
10:00am Randy Miller, Chestnut Hill College Tardigrades of the Sub-Antarctic: 5000 year old eggs from Marion Island
10:30am Morning Break Mediterranean Palm
11:00am Jesper Hansen, University of Copenhagen A Study on the Genus Amphibolus from Disko Island, Greenland, with Special Attention on the Life Cycle of Amphibolus nebulosus (Eutardigrada: Eohypsibiidae)
11:30am Matthew Boeckner, University of Newfoundland Factors Affecting the Ecology of Tardigrada in Labrador, Canada with Relation to Elevation, Latitude, Seasonality, and Desiccation Tolerance.
12:00 – 12:30pm Lunch Boxed Lunch
12:30pm Busch Gardens Resort Entrance
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 5
Date/Time Event Location______
Thursday July 31, 2003
Session 1 Life History -- Marine Mediterranean Palm
9:45am Tom Boesgaard, University of Copenhagen The Tardigrade Fauna of Two Australian Marine Caves With Descriptions of Six New Species of Arthrotardigrada.
9:45am Iben Heiner, University of Copenhagen A Revision of the Marine Genus Orzeliscus, Arthrotardigrada, Tardigrada
10:30am Morning Break Mediterranean Palm
11:00am Reinhardt Kristensen, University of Copenhagen Extreme Secondary Sexual Dimorphism in the Genus Florarctus (Arthrotardigrada: Halechiniscidae)
11:30am Clark Beasley, McMurry University Additions to the Tardigrada Fauna of China
12:00 – 1:30pm Lunch Royal Palm – 8th Floor
Session 2 Poster Session 2 Sabal - Canary Palm 3rd Floor
1:30pm Łukasz Kaczmarek, A Mickiewicz University Milnesium katarzynae sp. nov., a New Species of Eutardigrade from China
Hiroki Harada, Yokohama National University The response of Soil-Inhabiting Tardigrade Communities to Various Forests in the Southern Part of Kanagawa Prefacture.
Nigel Marley, University of Plymouth Tardigrades of Southwest England, United Kingdom. A Long-term, Multi-habitat Survey from the Coastal Urban Habitats to the Upland Moors.
Clayton Marshall, Eastern High School A Comparative Study of Souther Indiana Urban and Rural Tardigrade Populations Due to Seasonal Environmental pH Changes.
Harry Meyer, McNeese State University Small-scale Spatial Variability in Terrestrial Tardigrade Populations.
Maria Moly de Peluffo, Universidad Nacional de La Pampa Tardigrade Distributions in a Medium-sized City of Central Argentina
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 6
Date/Time Event Location______
Session 2 Poster Session 2- cont. Sabal - Canary Palm 3rd Floor
Lorena Rebecchi, University of Modena Resting Eggs in Tardigrades
Fran Thomas, University of Edinburgh Establishment of a Culture System for and Lifecycle Dynamics of the Tardigrade Hypsibius dujardini
Birna Trygvadottir, University of Copenhagen Tardigrades of the Faroe Islands
Karsten Klage, Virginia Tech University Tardigrades – Understanding Dessication Tolerance.
Sandra McInness, British AntarcticSurvey Tardigrade faunaof Sub-Antarctic Marion Island in the Prince Edward Archipelago, South Indian Ocean – A Preliminary Report.
Sandra McInness, British Antarctic Survey Tardigrade Fauna of the South Sandwich Islands
Randy Miller, Chestnut Hill College Tardigrades of North America: Central Park, New York City, New York, U.S.A.
3:00pm Afternoon Break Mediterranean Palm
Session 3 Taxonomy & Methods Mediterranean Palm
3:30pm Habib Maroon, University of Edinburgh Adult and Embryonic Anatomy of Hypsibius dujardini
4:00pm Randy Miller, Chestnut Hill College Auto-Montage Imaging for Tardigrades
4:30pm Jonnathan Herrera-Vásquez, University of Costa Rica Tardigrades Density and Diversity in Four Life Zones of Costa Rica.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 7
Date/Time Event Location______
Friday August 1, 2003
Session 1 Physiology Mediterranean Palm
9:00am Richard Helm, Virginia Tech University Tardigrades and Biomimetic Cell Stabilization
10:00am Lorena Rebecchi, University of Modena Long-term Anhydrobiotic Survival of Lichen-dwelling Tardigrades
10:30am Morning Break Mediterranean Palm
11:00am Roberto Guidetti, University of Modena Encystment in Eutardigrades: Differences and Common Traits in Two Evolutionary Lines
12:00pm Open Forum
1:00 – 2:00pm Lunch Royal Palm – 8th Floor
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 8
Semiterrestrial Tardigrades from Sakhalin Island, Far East Russia
Wataru ABE
Division of Biological Sciences, Graduate School of Science, Hokkaido University, Sapporo, Japan
Abstract. No report on the tardigrades from the Sakhalin Island has been made until now. In
July and August 2001 faunal survey on the semiterrestrial tardigrades was conducted in Sakhalin
Island, Far East Russia as a part of the International Sakhalin Island Project (ISIP), which is an international collaboration of Russia, USA, and Japanese scientists. Mosses and lichens growing mainly on trees and rocks were sampled to survey the tardigrade fauna. As a result, species belonging to the following 10 genera representing 4 families were collected: Echiniscus,
Hypechiniscus, Pseudechiniscus, Testechiniscus, Diphascon, Itaquascon, Ramazzottius,
Macrobiotus, Minibiotus, and Milnesium. Faunal characteristics of the tardigrades from the
Sakhalin Island will be discussed on the basis of the comparison with those from sorrounding areas including Japan.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 9
Phenotypic life history variations in two clones of Macrobiotus richtersi (Eutardigrada, Macrobiotidae)
Tiziana ALTIERO, Lorena REBECCHI, and Roberto BERTOLANI
Department of Animal Biology, University of Modena and Reggio Emilia, Modena, Italy.
Abstract. The study of life history traits is focal to understand animal evolution and adaptations.
Recent identification of rearing methods for freshwater and semiterrestrial eutardigrades under
controlled conditions (Altiero and Rebecchi, 2001, Zool. Anz. 240: 217-221), allowed us to overcome the limits imposed by the utilisation of animals exclusively derived from nature for the analysis of different aspects of tardigrade biology, including life history. Therefore, a comparative study of some life history traits has been realised under experimental conditions using two clones (namely, CDMr01 and CDMr02) of a triploid thelytokous apomictic population of Macrobiotus richtersi found in Italy. Both clones were reared at the same conditions: temperature of 14°C, photoperiod of 12 h/12 h (L/D), and nematode ad libitum as food. Intra- and interclonal variability has been observed for most life history traits analysed. Similarities between clones have been observed as regards life span, total number of ovipositions per life span, and age at first oviposition. The two clones were significantly different for number of eggs per clutch (fertility; p < 0.001), number of eggs laid by each female in its life span (fecundity; p
< 0.05), hatching percentage of eggs (p < 0.05) and embryonic development length (p < 0.001).
Having all specimens the same genotype, differences in life history traits between clones should be interpreted as phenotypic variations. Hatching phenology suggests that resting eggs could exist also in tardigrades, opening a new field of study on the life history traits of these animals.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 10
A Large-scale, Multi-Habitat Inventory of Tardigrades in the Great Smoky Mountains National Park
1 2 Paul J. BARTELS and Diane R. NELSON
1Department of Environmental Studies & the Environmental Leadership Center, Warren Wilson College, Asheville, North Carolina, U.S.A.
2Department of Biological Sciences, East Tennessee State University, Johnson City, Tennessee, U.S.A.
Abstract. An All Taxa Biodiversity Inventory (ATBI) is underway in the Great Smoky
Mountains National Park (GSMNP), attempting to identify all species of life in the 2000 km2 park. The GSMNP is a U.N. Biosphere Reserve and one of the largest protected temperate, deciduous forests in the world. We have completed two field seasons of work on the tardigrades of the park (2001-2002). To date we have collected over 400 samples from soil, lichen and moss on trees, stream sediment and periphyton, and caves. Terrestrial samples were taken from within permanent plots established for the ATBI, representing the major biological communities of the
GSMNP. Tardigrades were extracted from samples using centrifugation with LudoxTM,
individually mounted in Hoyer’s medium, and studied with phase-contrast microscopy. We have
identified 1663 specimens from approximately 70 samples. Only one study of tardigrades had
been previously reported for the park prior to our work, recording only three species. We have
identified 42 species, three of which may be new to science. Species richness estimates were
calculated for each of the major tardigrade habitats. Overall, our database predicts that there are
47 to 76 species in the GSMNP, with generally similar species richness in soil, lichen, moss, and
stream habitats. Species richness estimators were used to compare tree moss at ground level and
at breast height. Species richness was greater at breast height.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 11
Additions to the Tardigrada Fauna of China
1 2 3 CLARK W. BEASLEY , ŁUKASZ KACZMAREK , AND ŁUKASZ MICHALCZYK
1Department of Biology, McMurry University, Abilene, Texas 79697, U.S.A.; e-mail: [email protected]
2Department of Animal Taxonomy & Ecology, Institute of Environmental Biology, A. Mickiewicz University, Szamarzewskiego 91 a, 60-569 Pozna½, Poland; e-mail: [email protected]
3Institute of Environmental Sciences, Jagiellonian University, Ingardena 6, 30-060 Kraków, Poland; e-mail: [email protected]
Abstract. A total of 95 habitat samples, primarily lichen and moss, from the Chinese Provinces
of Sichuan and Yunnan were examined. The samples came from altitudes ranging from 2600 to
3850 m asl. Twenty-six species were recovered. One species, Milnesium katarzynae is new for
science and is described in a separate paper. Species which are new records for China include
Bryodelphax tatrensis, Echiniscus nepalensis, Echiniscus reticulatus, Echiniscus spiniger,
Isohypsibius sattleri, Diphascon (Diphascon) pingue, Diphascon (Adropion) scoticum,
Diphascon (Adropion) prorsirostre, Mesocrista spitsbergense, Platicrista angustata, and
Doryphoribius cf. zyxiglobus. Three species are new records for both Sichuan and Yunnan
Provinces: Macrobiotus cf. hufelandi, Minibiotus intermedius, and Hypsibius pallidus. Four
species are new records for Sichuan Province: Echiniscus sp. ‘arctomys-group’, Macrobiotus cf.
harmsworthi, Minibiotus sp., and Murrayon sp. Three species are new records for Yunnan
Province: Cornechiniscus lobatus, Pseudechiniscus jiroveci, and Doryphoribius citrinus.
Although not new records, Echiniscus testudo, Pseudechiniscus suillus, and Milnesium tardigradum were also collected from Sichuan Province. Bryodelphax tatrensis and Milnesium tardigradum were also found in moss samples from Xizang Province, Tibet.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 12
Phylogenetic relationships in Macrobiotidae (Tardigrada). II. Molecular (mtDNA) and morphological approaches
1 2 2 1 Roberto GUIDETTI , Andrea GANDOLFI , Valeria ROSSI , and Roberto BERTOLANI
1 Department of Animal Biology, University of Modena and Reggio Emilia, Modena , Italy
2 Department of Environmental Sciences , University of Parma , Parma , Italy .
Abstract. Molecular analyses on tardigrades are still at the beginning. First studies based on
18S rDNA considered the phylogenetic relationships between these organisms and other invertebrates, in particular arthropods and nematodes (Giribert et al. 1996; Garey et al. 1996;
Moon & Kim 1996; Aguinaldo et al. 1997). Successively, a molecular study (18S rRNA) within the phylum tested the validity of the actual classification at order level (based on morphological data; Garey et al. 1999). The present study has been carried out on Macrobiotidae (one of the most represented families of eutardigrades) using two different approaches: morphological and molecular. The morphological approach refers to up dated considerations presented in a previous paper (Guidetti & Bertolani 2001, Zool. Anz. 240: 371-376), whereas the molecular analysis has been done on mitochondrial DNA, which was considered for the first time in these organisms.
Eight species have been analysed at molecular level (Dactylobiotus parthenogeneticus,
Murrayon pullari, Macrobiotus terminalis, Macrobiotus sp., Macrobiotus richtersi, Xerobiotus pseudohufelandi, Ricthersius coronifer and Amphibolus volubilis as outgroup). The morphological and molecular data give similar results and confirm the previous phylogenetic evaluations on the main evolutionary lines (Murrayinae and Macrobiotinae) within the
Macrobiotidae. In particular, the strict phylogenetic relationship between Murrayon and
Dactylobiotus (Murrayinae) and the polyphyletic nature of Macrobiotus has been evidenced.
Macrobiotus sp. and M. terminalis (both belonging to the “hufelandi group”) look more related to X. pseudohufelandi than to M. richtersi. The R. coronifer position is still uncertain.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 13
Molecular phylogenetics of the Tardigrada and an investigation of the position of tardigrades in animal phylogeny
Mark BLAXTER , Ben ELSWORTH, Jennifer DAUB, Habib MAROON, Aziz ABOOBAKER, and Fran THOMAS
Institute of Cell, Animal and Population Biology, Ashworth Laboratories, University of Edinburgh, King’s Buildings, Edinburgh EH9 3JT, UK
Abstract: We have initiated a program of evolutionary developmental biology research on a
cultured tardigrade, Hypsibius dujardini. Tardigrades are an attractive organism for comparative
work because of their basal position in the pan-Arthropoda, and the observation that they share
some morphological and developmental characteristics with other Ecdysozoa such as the
nematodes. Many molecular and morphological analyses now agree that tardigrades should be
included within the Ecdysozoa, but have yielded divergent positions within the superphylum.
Small subunit ribosomal RNA genes place tardigrades in a nematode-priapulid branch, while
morphology unequivocally suggests an onychophoran-arthropod association.
We are investigating tardigrade phylogeny by (1) sequencing small subunit ribosomal RNA
genes from additional species and (2) using other nuclear and mitochondrial genes. We are able
to amplify SSU genes from individual tardigrades isolated from the wild and thus have
assembled a reasonable dataset of SSU sequences. The tardigrade-derived SSU dataset yields a
new view of tardigrade diversity. Our ongoing EST project on Hypsibius dujardini (see poster by
J. Daub et al) has yielded a near-complete set of mitochondrial genes, and also many nuclear genes that can be used to analyse tardigrade phylogeny. Our developmental biology program is also generating sequences for conserved regulatory genes such as HOX and PAR genes. The assembled datasets still yield a tardigrade-nematode association, but this appears to be due, to some extent, to long branch attraction.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 14
Factors Affecting the Ecology of Tardigrada in Labrador, Canada with Relation to Elevation, Latitude, Seasonality and Desiccation Tolerance.
1 1 2 Matthew J. BOECKNER , Michael A.J. COLLINS , and Lois BATEMAN
1 Department of Biology, Memorial University of Newfoundland , St. John’s , Newfoundland and Labrador , Canada
2 Science Division, Sir Wilfred Grenfell College , Memorial University of Newfoundland , Corner Brook , Newfoundland and Labrador , Canada
Abstract. Tardigrades are identified from mosses quantitatively sampled from varying
elevations, latitudes, seasons and horizon depths within coastal Labrador, Canada. Non-metric
multidimensional scaling is used to determine the relationships these environmental variables
have on the distribution, abundance and richness of tardigrade communities. Species that are
limited to specific environments as well as those with more cosmopolitan distributions are
indicated. Tardigrade distribution and richness throughout the samples was most strongly
correlated with the tendency for the moss to desiccate and most weakly correlated with latitude.
An elevational trend is discussed for M. c.f. hufelandi, which is strongly correlated with low
elevations. Nearly all of the tardigrade eggs were collected in the late summer and belonged to
M. echinogenitus, suggesting a strong relationship between reproduction and seasonality for this
species. Distribution patterns and ecological preference of individual species in relation to these
four environmental factors is discussed and compared to similar studies conducted worldwide.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 15
The Tardigrade Fauna of Two Australian Marine Caves . With descriptions of six new species of Arthrotardigrada
Tom M. BOESGAARD and Reinhardt Mbjerg KRISTENSEN
Invertebrate Department, Zoological Museum, University of Copenhagen , Denmark .
Abstract. Two marine caves in Australia have been investigated for meiofauna using the
freshwater technique to shock large sediment samples. Several species of nematodes, gastrotrichs, crustaceans, polychaetes and aplacophorans were found, and one new species of kinorhynchs has been described. Furthermore, two new species of loriciferans from marine caves in New South Wales , Australia are right now under description. This paper is the fourth in a
series describing the unique meiofauna in submarine caves and inland anchialine caves of
Australia . The paper give a full description of the tardigrade fauna of the caves, Jim's Cave and
Fish Rock Cave , both located off the coast of New South Wales . The sediment consist of carbonate sediments mixed with organic detritus.
The abundance of tardigrades is very low in the two caves, but the species diversity is very high. Until now the following arthrotardigrade genera are found: Actinarctus, Batillipes,
Dipodarctus, Halechiniscus, Raiarctus, Styraconyx, Tanarctus, Tholoarctus, and
Wingstrandarctus. Two species of Dipodarctus, two species of Wingstrandarctus, one species of
Batillipes, and one species of Tanarctus are new for science. The new species of Tanarctus is also
found in the North Atlantic . The cave fauna of tardigrades seems not to be very related to the well-
investigated high energy beach fauna of tardigrades along the East Coast of Australia. Most
surprising is the finding of several species known from the Italian caves (e.g. Arctinarctus
neretinus) supporting the theory that marine caves serves as refugees for an old Tethyan fauna.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 16
A Preliminary Account of the Tardigrades of Labrador, Canada
1 1 2 Michael A.J. COLLINS , Matthew J. BOECKNER , and Lois BATEMAN
1 Department of Biology, Memorial University of Newfoundland, St. John’s, Newfoundland and Labrador, Canada
2 Science Division, Sir Wilfred Grenfell College, Memorial University of Newfoundland, Corner Brook, Newfoundland and Labrador, Canada
Abstract. This is the first report of tardigrades in Labrador, the mainland part of the Canadian province of Newfoundland and Labrador. Seventeen species have been identified with a further three species yet to be identified. It had been expected that the tardigrades found in Labrador would overlap those of Greenland to the north, and of Newfoundland to the south, but the total species count is much more similar to the latter (n= 26) than the former (n= 80). Two species have been located in Labrador (Macrobiotus echinogenitus and Diphascon (Diphascon) recamieri) which have not been recorded for Newfoundland although both are found elsewhere in Canada. Proechiniscus hannae which had only been recorded for Western Greenland prior to the Newfoundland study has now also been recorded for Labrador. Diphascon ramazzottii which has previously been recorded for Europe and one location in Newfoundland was recorded in
Northern Labrador in very high numbers. Of the 31 species usually regarded as Arctic tundra species only 12 have so far been recorded for Labrador suggesting that the tardigrade fauna is somewhat depauperate for this region. Similarly of the seven heterotardigrades usually associated with Arctic environments only one has been found in Labrador.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 17
Origin and Diversification of the Arthropods: New Interpretations of Some Old Characters 1 2 Ruth Ann DEWEL and Jetta EIBYE-JACOBSEN
1 Department of Biology, Appalachian State University, Boone, North Carolina, U.S.A.
2 Invertebrate Department, Zoological Museum, University of Copenhagen, Copenhagen, Denmark
Abstract. The large morphological gaps separating extant taxa such as onychophorans, priapulids, and arthropods impede our ability to elucidate transitions that occurred in the early evolution of these organisms. The successful closing of these gaps will depend on finding fossil taxa, characters, and character states that can reveal critical steps in their evolution. However, the identification and characterization of potentially informative characters in fossil taxa depends in turn on understanding the structure, and if possible the development, of their putative homologues in living organisms. Several characters exhibit diverse character states in fossil and extant taxa and appear to be particularly useful in providing information on the early evolution of the arthropods. One such character is the buccopharyngeal apparatus of tardigrades, which is considered to have primary homology with portions of the “Peytoia” apparatus of Cambrian stem group arthropods. Taxa bearing a “Peytoia” are found among lobopods, stem group arthropods and euarthropods. The “Peytoia” apparatus is not considered to be homologous to the radially symmetrical mouth and introvert of cycloneuralians, but to be a circumoral novelty that incorporates at least one pair of limbs and surrounds a phylogenetically older mouth. The buccopharyngeal apparatus, together with characters such as the arthropod labrum and biramous limb, have been entered in a 44 taxa, 148 character phylogenetic analysis of primarily lobopod and arthropod ecdysozoans. Characters for this analysis were defined broadly to include previously unrecognized character states and have been described with explicit a priori statements of putative homology
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 18
Genomic resources for the tardigrade Hypsibius dujardini
Jennifer DAUB, Fran THOMAS, Habib MAROON, Aziz ABOOBAKER, and Mark BLAXTER
Institute of Cell, Animal and Population Biology, Ashworth Laboratories, University of Edinburgh, Kings Buildings, Edinburgh, EH9 3JT, UK.
Abstract: We have begun a programme of comparative developmental biology utilising the
cultured tardigrade Hypsibius dujardini as a new model for examining evolution of core
developmental processes. Tardigrades are an attractive organism for comparative work because
of their basal position in the pan-Arthropoda, and the observation that they share some
morphological and developmental characteristics with other Ecdysozoa such as the nematodes
and arthropods. This small fresh water species can be cultured in the laboratory on an alga food
source Chlamydomonas reinhardtii, and produces ample material for embryology and life cycle
studies (see poster by Thomas et al.).
As part of our programme of research we are developing genomic resources for H.
dujardini. We intend to construct and screen both genomic and cDNA libraries to isolate genes of
interest for use in phylogenetic and developmental studies. Thus far, a mixed stage cDNA library
has been constructed and screened by the expressed sequence tag strategy. So far over 900 ESTs
have been generated which represent ~500 individual tardigrade genes. The nuclear genes
identified include both housekeeping genes (encoding ribosomal proteins, cytoskeletal proteins ) as well as important regulator genes (encoding 14-3-3 proteins, Hox B4, kinases ). In addition, most of the mitochondrial protein- and rRNA-coding genes have been identified.
A preliminary analysis of the EST dataset will be presented. These ESTs will form the basis of subsequent in situ expression screens and phylogenetic analysis.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 19
The Ecological Distribution of Tardigrada in National Nature Reserve Stužica (Bukovské vrchy Mts , NE Slovakia )
Peter DEGMA and Miroslava PEČALKOVÁ
Department of Zoology, Comenius University , Bratislava , Slovak Republic
Abstract. Ecological distribution of terrestrial tardigrades was studied in National Nature
Reserve Stužica (Bukovské vrchy Mts, East Carpathians ). Values or categories of fourteen environmental variables were recorded during taking samples of mosses and lichens. Tardigrades extracted from the samples were mounted in Hoyer's medium and identified using phase microscopy. The data was statistically evaluated. A total of 4 780 specimens representing 33 tardigrade species (2 classes, 15 genera) were collected and identified from 150 samples. No significant differences were found in tardigrade abundance from samples collected at different
underbeds. No significant regression was found between tardigrade abundance and sample distance from upper substrate border. We came up with the same result of regression analysis
between the abundance of tardigrades and the position of mosses/lichens above ground level.
Only substrate thickness in case of presence and the sample distance from upper border of
substrate in case of quantity of different species were found as the significant gradient variables.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 20
Three dimensional understanding of the buccal apparatus of Echiniscus viridissimus (Peterfi, 1956) (Echinisciodea, Tardigrada)
1 2 Jette EIBYE-JACOBSEN and Ruth Ann DEWEL
1Invertebrate Department, Zoological Museum, University of Copenhagen, Copenhagen, Denmark
2Department of Biology, Appalacian State University, Boone, North Carolina, U.S.A.
Abstract. Using a combination of Transmission electron Microscopy and Scanning electron Microscopy a three dimensional understanding of the buccal cavity, the buccal tube and the stylets of Echiniscus viridissimus is presented. The foremost part of the buccal tube is continuous with the buccal cavity and the stylet sheaths and forms what looks like an arrowhead structure. The stylets when protruded out through the moth opening penetrate this structure and cross each other dorsoventrally. The buccal cavity at this level is not spherical but is an s-shaped opening giving room for the penetrating stylets and most probably guiding the direction of their movements. The buccal tube connects the arrowhead structure and the lumen of the pharynx. SEM preparations of the buccal tube from the related species Echiniscoides sigismundi (Schulze, 1865) reveal that the buccal tube is build from a great number of small spicule-like rods that are held together by an organic matrix in the functioning organ. These rods fall apart when the recently resynthesized organs in late simplex stage specimens are prepared. A close look at the TEM sections from different levels of the buccal tube of Echiniscus viridissimus reveals that the buccal tube of this species is also constructed from bundles of small spicule-like rods. SEM preparations from other related species reveal that the surface of the buccal tube of members of the genus Echiniscus is ridged. Information from TEM sections and the preparations of preactive stadium specimens indicates that the buccal tube is constructed from bundles of spicule-like rods throughout the entire genus. Additional evidence from TEM-sections of a simplex stadium specimen of Actinarctus doryphorus Schultz, 1935 suggests that this conclusion might possibly hold true for the entire class Heterotardigrada. The stylets are seen to be hollow in the SEM preparations. Throughout their entire length the stylets have a horseshoe shaped cross section that is open in the foremost part and forms a second groove in the posterior part. Implications of these observations on the function of the stylets are discussed.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 21
Population Dynamycs of Dactylobiotus grandipes Schuster et al., 1977 (Tardigrada) in a Neotropical Eutrophic Pond
1 1 1 María L. FERNÁNDEZ , Julio R. PELUFFO and María C. MOLY de PELUFFO
1Facultad de Ciencias Exactas y Naturales, Universidad Nacional de La Pampa, Uruguay 151, 6300 Santa Rosa, La Pampa, Argentina.
Abstract. This work is a population study of Dactylobiotus grandipes (Schuster et al., 1977)
during an annual cycle in a eutrophic limnotope of the Neotropical region, i.e. the “Laguna Don
Tomás” ((36º37’ S, 64º18’ O; 177 above sea level), in the city of Santa Rosa (La Pampa,
Argentina). The tardigrades related to the periphyton were studied by means of monthly
sampling of small underwater stalks between February 2001 and January 2002. Density was
expressed as individuals per cm3 of substrate. The nutritional status of the populations was identified, as well as the reproductive stage and molting condition. The size range of active individuals was measured, both in the simplex stage and in those with developing eggs. The population structure in terms of size class for each month was determined. The possible number of molts was estimated and the presence of precystic, cystic and free egg stages was recorded.
Active individuals were found only during the April to September period, with a population peak during May-June. D. grandipes exhibits seasonal trends in population dynamics, appearing in autumn and disappearing in spring. They seem to survive seasonal changes by remaining at low densities on the bottom and encysted on a substrate during the summer. Comparing the results obtained with those of populations of D.grandipes in Lake Tahoe in North America, a similar pattern of annual variation in water temperature can be observed. As to the individual size, the southern population shows lower minimal values at hatching and lower still respect of the first ecdysis, sexual maturity and maximum size. These differences could be related to differences in the vital cycle favorable period and to differences in water temperature.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 22
Encystment in eutardigrades: differences and common traits in two evolutionary lines
Roberto GUIDETTI, Deborah BOSCHINI, Lorena REBECCHI and Roberto BERTOLANI
Department of Animal Biology, University of Modena and Reggio Emilia, Modena, Italy.
Abstract. Tardigrades have two forms of dormancy: cryptobiosis and encystment. The encystment is a form of diapause known for a limited number of tardigrades and it is still little studied. To increase the knowledge on encystment, two species of eutardigrades from Italy have been considered: the moss-dwelling Amphibolus volubilis (Eohypsibiidae), able both to enter cryptobiosis and to form cysts, and the freshwater Dactylobiotus parthenogeneticus
(Macrobiotidae), only able to form cysts. Cysts have been collected in nature or have been induced under laboratory conditions. In the latter case, it was possible to follow the encystment process phases. Cyst morphology has been analysed by LM, SEM and TEM. Two different types of cyst have been found in A. volubilis, while in D. parthenogeneticus only one type. In all three kinds of cyst, the encystment processes show both common and peculiar traits. Encystment begins with the discharging of the sclerified parts of the buccal-pharyngeal apparatus, as in ecdysis, but without the loss of the old animal cuticle. Then, two or three new cuticles are serially synthesized, according to the type of cyst. In A. volubilis, the ultrastructure of these new cuticles is similar to the active and unencysted animal cuticles while in D. parthenogeneticus the new cuticles ultrastructure differs from that of the active and unencysted animals. A modified buccal-pharyngeal apparatus, up to date undescribed, has been observed in one type of A. volubilis cyst and in the D. parthenogeneticus cyst. The common traits lead us to suppose a common origin of the phenomenon. These peculiarities may represent diversified adaptation strategies to different environments which should be studied more in depth.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 23
Dactylobiotus octavi n. sp. (Eutardigrada; Macrobiotidae) from Disko Island (Greenland)
Roberto GUIDETTI, Tiziana ALTIERO, and Roberto BERTOLANI Department of Animal Biology, University of Modena and Reggio Emilia, Modena, Italy.
Abstract. During the "Workshop on Arctic tardigrades" at the Danish Arctic Station
(Qeqertarsuaq, Disko Island, Greenland) organized by R.M. Kristensen and his co-workers at the
end of the VIII International Symposium on Tardigrada (Copenhagen 2000), an undescribed
species of Dactylobiotus was found in freshwater sediments of the creek Isunngua. We had the
honour and the pleasure to describe this new taxon that we would like to dedicate to all
participants of that symposium, naming the species Dactylobiotus octavi n. sp. The animals look
similar to Dactylobiotus dispar and Dactylobiotus haplonyx for the presence of a very short secondary branch in the claws of their first three pairs of legs, but they differ from these species
for their claw size and for their buccal tube width. This new species also has peculiar ornamented
eggs. The egg shell consists of open, crater-like processes connected up to the apex. This finding
increases the already high number of species found in Disko Island and once again underlines the
tardigrade importance in the biodiversity not only in that island, but in all the Arctic area. It also
confirms the peculiarity of some Dactylobiotus characters, which represents a very well defined
evolutionary line of macrobiotids developed in freshwater.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 24
The “Hyena Female” within the Marine Tardigrada with the Description of Two New Species of Megastygarctides (Arthrotardigrada: Stygarctidae) from Saudi Arabia.
Jesper Guldberg HANSEN and Reinhardt Møbjerg KRISTENSEN
Invertebrate Department, Zoological Museum, University of Copenhagen, Universitetsparken 15, DK-2100 Copenhagen Ø, Denmark
Abstract. The Arabian Gulf of Saudi Arabia has a rich tardigrade fauna. The authors have
studied 22 small vials with tardigrades from the ARAMCO Northern Area Intertidal Sampling
Program in 1982. Eleven arthrotardigrades and one echiniscoid species were recorded from
seven intertidal stations. In this paper we described two new species of Megastygarctides. Slight secondary sexual dimorphism is present in all species of the family Stygarctidae, but in one of the new species, both the primary and secondary clavae have a different shape in the male than in the female. Furthermore, the female has a unique genital structure. The genital ducts of the two seminal receptacles are extended out of the body as two robust penile spines. The function of these two structures, located lateral to the rosette-shaped female gonopore, is still mysterious, but they may be involved in both copulation and in the insemination of the spermatozoa through the eggshell. The name suggested for this type of female in tardigrades is the “tardigrade hyena female”. The male has a typical heterotardigrade gonopore consisting of a small oval papilla with a crescent-shaped opening. Another unique character of this species is the plate pattern.
Typically, the plates are dorsal in the Stygarctidae but in this species the plates completely encircle the body. The other new species of Megastygarctides completely lacks dorsal plates, which is very atypical in the family. The two new species are compared with the known species of Megastygarctides and a revision of the genus is given.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 25
A Study of the Genus Amphibolus from Disko Island, Greenland, with Special Attention on the Life Cycle of Amphibolus nebulosus (Eutardigrada: Eohypsibiidae)
1 2 1 Jesper Guldberg HANSEN , Agnete Krabbe KATHOLM and Reinhardt Møbjerg KRISTENSEN
1Invertebrate Department, Zoological Museum, University of Copenhagen, Universitetsparken 15, DK-2100 Copenhagen Ø, Denmark
2 Department of Life Science and Chemistry, Roskilde University, Roskilde, Denmark
Abstract: In the last thirty years numerous investigations of the Disko Island tardigrade fauna has been carried out from the Danish Arctic Station, Qeqertarsuaq, Disko Island, West Greenland. The results have partly been obtained by scientific leaders of the Arctic Station and partly from several field courses in Arctic Biology. At present, more than a hundred different species of limnic/terrestrial tardigrades have been collected on the Disko Island. As part of a field course in Arctic Biology, samples of mosses from four habitats were collected on Disko Island, Greenland, in July 2002, in order to study the life cycle of Amphibolus nebulosus. Additional material collected on Disko Island in the period 1976-2001 was examined for the occurrence of Amphibolus species. The three species A. nebulosus, A. weglarskae and Amphibolus nov. sp., are unequally distributed on the Disko Island at 24 locations. The environmental preferences of A. nebulosus are apparently for wet habitats, as running or calm water, for mud, soil and mosses, and for more dry environments as lichens. Based on these data, it seems probable to consider A. nebulosus being hygrophilous and not a true hydrophilous species. The environments preferred by A. weglarskae are drier (e.g. soil) than those preferred by A. nebulosus. The two species are hardly ever found in the same substrate. Living in a range of dry to moist habitats, it seems that A. weglarskae is a eurytopic species. Amphibolus nov. sp. is probably a real hydrophilous tardigrade. It is found in permanent freshwater habitats like springs, lakes and rivers. We found Amphibolus nov. sp. in aquatic mosses and algae in a heterothermic pool, where we also found A. nebulosus. The study of Amphibolus nebulosus signifies that the life cycle involves two types of cysts and two types of eggs. It seems that both kinds of cysts are related to reproduction as well as to environmental changes. New information on the sclerified structures, claws and the characteristics of the egg-shell within the genus are presented, and a modification of terminology is suggested.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 26
The response of soil-inhabiting tardigrade communities to various forests in the southern part of Kanagawa Prefecture
1 1 Hiroki HARADA & Masamichi T. ITO
1Soil Ecology Laboratory, Graduate School of Environment and Information Sciences, Yokohama National University, Yokohama-city, Kanagawa, Japan
Abstract. This study was carried out for the purpose of detecting the relationships between soil- inhabiting tardigrade communities and various types of forest. Nine sites were selected in the southern part of Kanagawa Prefecture. Vegetation types of these forests were: evergreen broad- leaved forests (4 sites), deciduous broad-leaved forests (1 site), evergreen coniferous forests (3 sites) and citrus fruit orchard (1 site). In these sites, environmental factors (e.g. leaf litter dry weight, soil pH, soil hardness, soil moisture content) were measured. For collecting tardigrades, Baermann method was adopted. Tardigrades were identified in specific level by using a DIC microscopy. Statistical analyses and multivariate analyses (CCA) were driven to know 1) species diversity, 2) community similarity and 3) the correlation between environmental factors (23 series) and tardigrade faunal composition. Tardigrade fauna was different among every forest sites, highest abundance (18,650 ind./m2) and largest species number (26 species) was occurred in an artificial coniferous forest (site “Nebu”). Through this study, two main groups of tardigrades were distinguished. First group (M-group) majorly contained Macrobiotus species, which is known as cosmopolitic species. Second group (D-group) was formationed by genus Diphascon (e.g. D. nobilei, D. patanei, D. prorsirostre). As remarkable fact, only Diphascon pingue was included to M group. D-group was concentrated in the site “Nebu”. On the contrary, M group species were dominant in other sites. From the result of CCA, distinct environmental factor could not be decided, but the frequency of nematodes was recognized as main factor which influences D- group existence. Particularity of the site “Nebu” was also proved by statistical data, and this result doesn’t correspond with large-scaled vegetational classification. It is sure that Nebu’s coniferous forest created the unique environment for sustaining these special species (D-group). As a conclusion, the forests should be evaluated not only by macroscopic factor, such as landscape, but by microscopic organisms, such as tardigrade communities.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 27
A Revision of the Marine Genus Orzeliscus, Arthrotardigrada, Tardigrada
Iben HEINER and Reinhardt M. KRISTENSEN
Department of Invertebrate Zoology, Zoological Museum of Copenhagen, Denmark, e-mail: [email protected] & [email protected]
Abstract. Du Bois-Reymond Marcus described the marine genus Orzeliscus with the species
belopus in 1952 from the island of São Sebastião in Brazil. In 1953 Schulz described
septentrionalis, which was synonymized with belopus in 1980 by Pollock.
Over the years different scientists have reported several findings of the genus Orzeliscus from
several other countries, e.g., France, New Caledonia, Bermuda, Scotland, the Virgin Islands and
the Galapagos Islands. The large collections in the Zoological Museum of Copenhagen,
Denmark include several specimens from France, Egypt, Japan, Bermuda, Tobago, USA and
Australia. Close examination of them has revealed the presence of several new species, e.g., a
new species from Japan with a protruding mouth cone and one from Egypt which is
hermaphroditic. The characteristics of these new species will be presented together with a
revision of the genus. The specimens from Queensland, Australia consist of three species, one of
which has only three toes on the each leg and lateral projections on the body with long pillars.
On the basis of these new characters a new genus has been established and the family
Orzeliscidae is rediagnosed. A map of the world distribution of the family with all known
records is also included.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 28
Tardigrades and Biomimetic Cell Stabilization
Richard F. HELM, Karsten KLAGE and Malcolm POTTS
Department of Biochemistry and Virginia Tech Center for Genomics, Virginia Tech, Blacksburg, Virginia USA
Abstract. The ability of tardigrades to withstand a wide variety of environmental insults has been a source of awe as well as a direction of scientific inquiry since their initial description by
Van Leeuwenhoek in 1702. As they are considered one of the most resilient animals on the planet, we are interested in understanding their metabolic biochemistry and cell biology with the long term vision of applying similar strategies to the preservation of cells and cell components of biomedical importance. Our present efforts focus on strategies to monitor the 3 major “-omes,” namely the transcriptome, proteome and metabolome as tardigrades undergo desiccation and rehydration. An overview of our research goals will be presented from the perspective of understanding the biochemical control of cryptobiosis and its effects on cell biology and metabolism.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 29
Seasonal and Spatial Variation in Tardigrade Diversity in Leaf Litter from Florida and Louisiana
Juliana HINTON, Harry A. MEYER, Kathleen TRAHAN
Department of Biological and Environmental Sciences, McNeese State University, Lake Charles, Louisiana, U.S.A.
Abstract. During 2002-2003, we collected leaf litter core samples (10cm diameter) from each of two sites in southwestern Louisiana and one site in central Florida. A mixture of deciduous leaves and pine needles characterized the Florida site and one of the Louisiana sites; the leaf litter of the other Louisiana site was composed almost entirely of needles. In Louisiana, cores were collected at four times: summer, fall, winter, and spring. Florida cores were only collected in winter and spring. Four cores were collected for each combination of site and date. Each core was divided into two layers, an upper leafy layer (1-2cm in depth), and a lower layer of humus
(1-2cm in depth). Five species of tardigrade were found in Louisiana material and three in
Florida. Within each site, there was wide variation in tardigrade diversity among samples and dates. Among the three sites, tardigrade species richness and abundance declined with the proportion of the material made up of needles. Tardigrade diversity at these sites in Florida and
Louisiana is considerably lower than that previously reported in leaf litter and humus samples from beech forests in Italy and Tennessee.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 30
The Effects of Prehydration on Anhydrobiotic Survival in the Tardigrade Milnesium tardigradum
1 2 1 Daiki D. HORIKAWA , Wataru ABE , and Seigo HIGASHI
1Division of Biosciences, Graduate School of Environmental Earth Science, Hokkaido University, Sapporo, Japan.
2Division of Biological Sciences, Graduate School of Science, Hokkaido University, Sapporo, Japan
Abstract. It has been thought that the high humidity prior to rehydration with water ensures the revival of anhydrobiotic animals. For example, it is known that placing anhydrobiotic nematodes directly in water at lower relative humidity causes decreased survival. However, there are little reports on the effects of prehydration on tardigrades. In the present study we estimated the effects of relative humidities on the recovery of the tardigrade Milnesium tardigradum from its anhydrobiotic state. The tardigrades were dried at 80% RH for 24 hours, and transferred to 0%
RH air for 72 hours before prehydration at various relative humidities (0, 25, 84 or 97%) for 24 hours. Then animals were rehydrated with water and showed high survival rates under all the conditions, indicating that this xerophilous tardigrade species does not require any prehydration when the animals revive from anhydrobiosis and is more tolerant to the rapid rehydration than other anhydrobiotic animals.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 31
Tardigrades in six phorophytes of the moss Calymperes tenerum C. Müller
1 Javier JEREZ - JAIMES
1Department of Biology, University of Puerto Rico, Mayagüez, Puerto Rico, U.S.A.
Abstract. The composition of tardigrade communities on the Mayagüez Campus of the
University of Puerto Rico was studied from January through May 2003. Eleven trees of six
phorophytes species were selected for the moss Calymperes tenerum: (Calophyllum brasiliense
(2), Swietenia macrophylla (2),Bucida buceras (2),Hymenaea courbaril(1), Mangifera indica (2) and Ptychosperma elegans (2). During the dry season (January- March) and the rainy season
(April-May) five samples of four cm2 each were taken from each tree. Tardigrades were
extracted from the samples, mounted individually in Hoyer’s medium, and identified to species
using phase and Nomarsky microscopy. Differences in tardigrade community composition were
founded among and within phorophyte species. Swietenia macrophylla and Mangifera indica were the phorophytes of the moss C. tenerum with the greatest number of species and abundance of tardigrades.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 32
Milnesium katarzynae sp. nov., a new species of eutardigrade (Milnesiidae) from China
1 2 3 Łukasz KACZMAREK , Łukasz MICHALCZYK and Clark W. BEASLEY
1Department of Animal Taxonomy & Ecology, Institute of Environmental Biology, A. Mickiewicz University, Szamarzewskiego 91 a, 60-569 Poznań, Poland; e-mail: [email protected].
2Institute of Environmental Sciences, Jagiellonian University, Ingardena 6, 30-060 Kraków, Poland; e-mail: [email protected].
3Department of Biology, McMurry University, Abilene, Texas 79697, U.S.A; e-mail: [email protected].
Abstract. A new eutardigrade, Milnesium katarzynae sp. nov. is described from two moss samples collected by Katarzyna Ratyńska in China (Nature Reserve near Kangding) in August
2002. Until now only 5 species of the genus Milnesium Doyere are known: Milnesium brachyungue Binda & Pilato, Milnesium eurystomum Maucci, Milnesium slovenskyi Bertolani &
Grimaldi (known only from Cambrian amber), Milnesium tardigradum Doyére and Milnesium tetralamellatum Pilato & Binda. This new species differs from other described members of genus in having fine (0.5 – 1.0 µm) reticular design on the dorsal side of the body (better visible in the caudal region), lacking of eyes, and having claws in the almost same size on all legs. M. katarzynae sp. nov. is similar to specimens of M. tardigradum from New Zealand which have reticular “shallow depressions” in cuticle but is clearly different by lacking of eyes and narrower buccal tube (pt = 24.8-29.6 in new species and pt = 41.9 in the specimens collected in New
Zealand).
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 33
Tardigrades – understanding desiccation tolerance
Karsten KLAGE1, Richard F. HELM1, Jonathan D. EISENBACK2, Ruth DEWEL3, Roberto BERTOLANI4 and Malcolm POTTS1
1Department of Biochemistry and Virginia Tech Center for Genomics, Virginia Tech, Blacksburg , Virginia 24061
2Department of Plant Pathology, Physiology and Weed Science, Virginia Tech, Blacksburg , Virginia 24061 USA
3 Department of Biology, Appalachian State University , Boone , North Carolina 28608 USA
4 Department of Animal Biology, University of Modena and Reggio Emilia, 41100 Modena , Italy
Abstract. Some species of tardigrades have the ability to survive long periods of complete dryness assuming a physiological state known as ‘tun’. It was shown that in this dry stage the tardigrades are able to resist diverse and extreme environmental conditions, such as low and high temperature (-195°C to 110°C), ionizing radiation, vacuum, high pressure (6000 times atmospheric) and long periods of dryness (up to eight years). However, by adding water to the dried organisms, they regain normal vitality with apparently no damage to cell organelles or cell structure. How can proteins, DNA, lipids, sugars, etc. withstand this water loss, without precipitating or denaturing, and what are the mechanisms that permit the tardigrade to survive complete dryness? Early work with tardigrades, and other desiccation tolerant organisms such as the brine-shrimp (e.g cyst of Artemia salina), lead to the development of the “water replacement hypothesis,” which describes how the non-reducing disaccharides, trehalose and sucrose, replace hydrating water molecules lost during desiccation. Later the hypothesis was modified through incorporation of the “glassy state theory”. Sugars and proteins like LEA create a ‘glass’ during dryness and provide stability and help maintaining cell integrity. Also emphasis was put on the potential role of amphiphiles. Using molecular and biochemical methods our laboratory is focusing on the mechanisms used by tardigrades to achieve desiccation tolerance. These include differential expression of mRNA, proteome analysis through mass spectrometry, electron microscopy, and analysis of sugars and lipids.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 34
Extreme Secondary Sexual Dimorphism in The Genus Florarctus (Arthrotardigrada: Halechiniscidae)
Reinhardt Møbjerg KRISTENSEN
Invertebrate Department, Zoological Museum, University of Copenhagen, Universitetsparken 15, DK-2100 Copenhagen Ø, Denmark
Abstract. Secondary sexual dimorphism in florarctid tardigrades is well known. The males are usually smaller than the female, and the primary clavae are relative longer in the males. A new species of Florarctus from coralline sand collected subtidally just behind the reef fringe of Long Island, Chesterfield Reefs (Pacific Ocean) has extreme secondary dimorphism. The male has developed clavae that are much thicker and three times longer than in the female. Furthermore, the clava of the male is formed as an accordion-like structure. This structure is not seen in the female where the clava is smooth. Ten other species of Florarctus were investigated from the Pacific Ocean. The male was always smaller than the female, but the male of the F. heimi and the female of F. cervinus have not been recorded. The description of F. heimi date back to 1965 and that of F. cervinus is from 1987. Renaud-Mornant described these two species from New Caledonia and her large collection was reinvestigated in the present study. In the Australian summer (December 1995) large populations of Florarctus species were found subtidally in Shark Bay, Heron Island, and the Great Barrier Reef. F. heimi and F. cervinus were found together in coralline sand from Heron Island. The animals were kept alive and video-taped in the laboratory of Queensland Museum. All specimens of the very large F. heimi (about 400 µm) were females and all specimens of the smaller F. cervinus (about 170 µm) were males. The differences in the caudal expansion between the two “species” were exactly as in the two excellent original descriptions. Males of F. cervinus were observed to mate with females of F. heimi. Observations of mating in Arthrotardigrada are rare. Only one observation of Parastygarctus sterreri has been made. As in P. sterreri the floractids mate venter to venter. The conclusion based on the observations on animals from Heron Island is that F. cervinus is a junior synonym of F. heimi. Florarctus cervinus is the male of F. heimi and this is the first time that such an extreme sexual dimorphism has been observed in tardigrades.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 35
Preliminary Results from a Study on Ecuadorian Tardigrada.
Nigel J. MARLEY
School of Biological Sciences,University of Plymouth, Drake Circus, Plymouth, PL4 8AA,United Kingdom .
Abstract. There are very few published studies on the tardigrade fauna of Ecuador. Preliminary results from an altitudinal survey of moss and lichen inhabiting tardigrades from the Volcán
Chiles, Ecuador are presented.
Sixteen operational taxonomic units have been found so far. Precise identifications, particularly for eutardigrades, has been difficult due to problems accessing literature on South American taxa. One species new to science is described, Platicrista ramsayi.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 36
Designation of Pseudobiotus kathmanae Nelson (Tardigrada) as the type species of Pseudobiotus Nelson.
1 2 3 Nigel J. MARLEY , Roberto BERTOLANI & Diane R. NELSON
1 School of Biological Sciences,University of Plymouth, Drake Circus, Plymouth, PL4 8AA , United Kingdom .
2 Dipartimento di Biologia Animale, Università di Modena e Reggio Emilia, Via Campi 213/d, 41100 Modena, Italy.
3 Department of Biological Sciences, East Tennessee State University, Johnson City, TN 37614 ,USA
Abstract. The contents of the application to the International Commission on Zoological
Nomenclature, Case 3017, are presented with an explanation of how its publication fell between the third and fourth editions of the Code of Zoological Nomenclature. The aim of the application was to ask the Commission to designate a new type species for Pseudobiotus Nelson, following the discovery of the misidentification of the original type. The full publication of case 3017 was then delayed until all papers relevant to the application were published: Nelson, Marley and
Bertolani (1999) and Bertolani, Marley and Nelson (1999). These two papers were also presented at the 7th International Symposium of the Tardigrada held in Düsseldorf, Germany. The case was then held by the Commission pending the publication of the fourth edition of the Code because of relevant changes to Article 70. The fourth edition of the Code came into affect on January 1st
2000 . Case 3017 then became unnecessary for the Commission to see and was left unpublished and without an Opinion ruling from the Commission. This has resulted in Pseudobiotus Nelson having no designated type. To rectify this situation the details of Case 3017 are briefly present here and then, with the reference to the appropriate articles of the fourth edition of the code, a new type is designated for the genus.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 37
Tardigrades of Southwest England, United Kingdom. A long term, multi- habitat survey from the coastal urban habitats to the upland Moors. (Short title: Bear Hunting in Devon)
Nigel J. MARLEY
School of Biological Sciences, University of Plymouth, Drake Circus, Plymouth, PL4 8AA , United Kingdom .
Abstract. The tardigrade fauna of southwest England has been poorly reported. A long term, low intensity survey has been carried out across the region. Samples have covered a wide range of habitats including rocky intertidal, deciduous woodlands, acid moorlands and urban xeric.
Sample materials have included seaweed, marine sediments, barnacles, bryophytes, lichens, leaf litter and bark. The results from this wide ranging survey are presented with many new additions to the local and national fauna being reported. One species new to science is described from the coastal urban habitats. The distribution of individual species and different tardigrade communities is discussed and related to bedrock type and water regime.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 38
Adult and Embryonic Anatomy of Hypsibius dujardini
Habib MAROON, Aziz ABOOBAKER, Fran THOMAS, Jennifer DAUB, and Mark BLAXTER
Institute of Cell, Animal and Population Biology, Ashworth Laboratories, University of Edinburgh, Kings Buildings, Edinburgh, EH9 3JT, UK.
Abstract. To gain a deeper understanding of the evolution of the Ecdysozoa (a superphletic grouping of moulting animals including nematodes and arthropods), we have recently initiated research into tardigrade biology using a a cultured Eutardigrade: Hypsibius dujardini (see abstract by Thomas et al.). We aim to use this species to study tardigrade development to further our understanding of the evolution of developmental processes that have been studied deeply in arthropod and nematode model systems. Here we describe progress on the characterisation of the embryonic and adult anatomy of Hypsibius dujardini. We are developing methods to introduce vital and other stains into live and fixed animals. Currently we are able to stain juveniles and adults reliably, but embryos are relatively diffcult to stain (presumably because of permeability barriers in the egg). DAPI staining of nuclear DNA reveals cell number in the different organ systems of adults. Phalloidin, a stain that binds to actin cables, displays the musculature of the adult. Bodipy-Fl-Ceramide, a membrane-associated fluorescent marker, has been used to show the nervous system. We have also undertaken light and fluorescent microscopic studies of embryonic development confirming some previous observations (Eibye-Jacobsen.1997). DAPI staining has also been of some use for investigating embryonic development. We are currently endeavouring to define suitable protocols for permeabilisation of the chorion, immunohistochemistry, and for in situ hybridisation.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 39
A Comparative Study of Southern Indiana Urban and Rural Tardigrade Populations Due to Seasonal Environmental pH Changes
Clayton MARSHALL
Eastern High School, Pekin, IN
Abstract. Tardigrades are invertebrates that are used as an environmental bio-indicator.
Problems with acid precipitation are occurring across America’s Northeast region. Coal burning
plants along the Ohio River Valley are a suspected cause of these problems. The purpose of this
research project was to determine the effect of environmental pH on tardigrade populations collected from lichen samples on urban and rural limestone monuments in Southern Indiana. It was hypothesized there would be a positive correlation between environmental pH and tardigrade community structure. Testing sites were selected from six urban and six rural areas in
Southern Indiana, and four tests were conducted during each of the four seasons resulting in a total of 96 samples collected. Lichen samples were placed into distilled water, and tardigrades were extracted, counted, and preserved. The pH of each sample was then measured. Tardigrades were later mounted on glass slides and were identified as the following species: Milnesium tardigradium, Echiniscus perviridis, Echiniscus viridis, Echiniscus cavagnaroi, and Echiniscus knowltoni. The 12-month urban pH mean was 7.25 with a total tardigrade population of 96. The
12-month rural pH mean was 7.07 with a total tardigrade population of 1292. The deviation of pH from neutral was statistically analyzed using a t-test and was determined to be highly significant at the 4 x 10-6 level. The correlation between pH and tardigrade population density was statistically analyzed using linear regression and was determined to be significant at the 0.01 level. Based on the data collected the hypothesis was supported.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 40
Exceptional, Tardigrade Dominated Ecosystems in Ellsworth Land, Antarctica
Sandra J. MCINNES and Peter CONVEY
British Antarctic Survey, Natural Environment Research Council, High Cross, Madingley Road, Cambridge CB3 0ET, UK
Abstract. We describe an exceptionally simple terrestrial faunal community present on inland nunataks of Ellsworth Land, Antarctica, at locations between Sky Hi and Haag Nunataks (c. 75 -
77°S 70 - 73ºW). No biological studies have previously been made at these latitudes in the
Antarctic Peninsula (West Antarctic) region. Samples obtained from a series of isolated mountain groups indicate a common fauna dominated by Tardigrada, with a minority component of Rotifera. The fauna is exceptional in its simplicity, including 6 tardigrade species (1-2 new to science) and two trophic levels. Neither nematode worms, the most important element of the simplest communities previously reported worldwide (from the Ross Sea Dry Valley region of continental Antarctica), nor microarthropods, otherwise represented in all known Antarctic terrestrial communities, are present. The community shows closer affinities with continental
Antarctic tardigrade communities, with which it shares three species, than the maritime zone, sharing only two pan-Antarctic species with the latter. The remaining four species form a group that is unique to Ellsworth Land, and may suggest its prolonged existence as a distinct biogeographical unit.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 41
Tardigrade fauna of Sub-Antarctic Marion Island in the Prince Edward Archipelago, South Indian Ocean – a Preliminary Report.
Sandra J. MCINNES
British Antarctic Survey, Natural Environment Research Council, High Cross, Madingley Road, Cambridge CB3 0ET, UK
Abstract. Sub-Antarctic Marion Island (46°54’S; 37°45’E) one of the islands in the Prince
Edward Archipelago, South Indian Ocean, is a c. 4 million years old shield volcano, with the remnants of a summit Pleistocene glacial ice cap. A survey was carried out in April 2003 to identify the tardigrade species diversity and habitat preferences of the fauna on this island.
Limno-terrestrial tardigrades were sampled from a range of terrestrial habitats that included moss, lichen, soil and cushion forming higher plants. Freshwater habitats were sampled from shallow shelf regions and marine tardigrades collected from the coastal shorelines. Collections focused on specific terrestrial habitats and produced several new records for the island and new species.
The tardigrade records from this study represent the first detailed reports from Marion Island, and are to be compiled into a database with three primary objectives; 1) elucidate the biodiversity of sub-Antarctic Marion Island tardigrades; 2) identify ecological, in particular substrate association, of the limno-terrestrial fauna; and 3) associations with other co-occurring meiofauna.
The limited earlier references to tardigrades on Marion Island placed the island group in the biogeographic sub-Antarctic cluster. This new study adds more detail to the biogeography of the tardigrades in this sector, and forms one of the baseline studies on the meiofauna of Marion
Island. St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 42
Tardigrade fauna of the South Sandwich Islands
Sandra J. MCINNES and Peter CONVEY
British Antarctic Survey, Natural Environment Research Council, High Cross, Madingley Road, Cambridge CB3 0ET, UK
Abstract. Sub-Antarctic South Sandwich Islands comprise a group of islands of volcanic origin between the latitudes of 56º 18'S, 27º 34'W and 59º 27'S, 27º 22'W. All the islands are of recent origin with many still exhibiting some form of volcanic activity from warm ground and hot lakes to smoking fumaroles. The islands are part of a crustal upwarp extending from South Georgia through the South Sandwich Islands to the South Shetland Islands that connects the Andean chain of South America to the Antarctic Peninsula.
These islands were part of an extensive survey during 1997, covering vegetation and meiofauna.
The tardigrade fauna shows a limited biodiversity, in keeping with the recent formation of these islands. However, there is an indication that the source of the fauna is from both sub-Antarctic and Antarctic origins.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 43
Distribution of Terrestrial Tardigrades in the State of Florida
Harry A. MEYER
Department of Biological and Environmental Sciences, McNeese State Universit , Lake Charle , Louisiana, U.S.A
Abstract. The distribution of terrestrial tardigrades in the state of Florida is poorly known.
Published records from the state include only two species, Echiniscus perarmatus and E. virginicus, both from the northeastern part of the state. I collected samples of moss, lichen, and liverwort from all 67 Florida counties during 2002-2003. These samples were primarily taken from trees and fallen branches. Where possible, I chose sampling sites in the relatively undisturbed wooded areas of state and national parks. However, a large portion of the Florida landscape has been heavily impacted by agriculture, forestry, and urban development. Therefore, the sampling sites in many counties were in highly disturbed habitats (rural roadsides, cemeteries, municipal parks, urban neighborhoods, etc.). Where possible, lichens, mosses, and liverworts were identified (27 species). The number of tardigrade species per sample ranged from one to four. In all, sixteen tardigrade species, including E. virginicus, were found in the state. The most commonly detected species were Milnesium tardigradum, Macrobiotus echinogenitus, and Minibiotus intermedius. Echinscids and non-Macrobiotid species of the Order
Parachela were relatively uncommon. Suitable tardigrade habitat was difficult to locate in much of the central and southern regions of the state, especially in pine forests, the Everglades , and the
Florida Keys . Tardigrade species richness and abundance also appeared to be lower in these areas. Preliminary results from a similar, as yet uncompleted, survey of Louisiana suggest that tardigrade diversity may be lower in Florida than in Louisiana.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 44
Small-scale Spatial Variability in Terrestrial Tardigrade Populations
Harry A. MEYER
Department of Biological and Environmental Sciences, McNeese State University , Lake Charles , Louisiana , U.S.A.
Abstract. Terrestrial tardigrades commonly occur in the lichens and mosses that occur on trees
and rocks. Although tardigrades in these habitats are often said to be very patchy in their distribution, this assessment has not often been backed by quantitative sampling. In this study I assess spatial variability in tardigrade populations inhabiting small patches (1-45mm diameter) of moss and lichen on trees and rocks. In 2002 I collected tardigrades from four replicate rocks in the Ouachita Mountains of Arkansas, U.S.A. I collected 30 lichen patches on two rocks and 20 moss patches on two others. In Lee County, Florida I collected tardigrades from lichen patches on two neighboring Royal Palm Trees. The tardigrades in each sample were mounted and identified and the numbers of bdelloid rotifers, nematodes, and mites recorded. The variation among lichen or moss patches within rocks or trees was very high; the only consistent patterns were that very small patches usually lacked tardigrades, and the predatory species Milnesium
tardigradum tended to be most abundant in larger patches. Tardigrade diversity abundance also
varied greatly within sites when lichens and mosses of the same species were compared from
different rocks and trees (in the most extreme case, comparing the two Royal Palms, one tree had numerous individuals of three tardigrade species present while the other had no tardigrades at all). The results of this quantitative sampling support the assertion that tardigrades are very patchy in distribution. Given the considerable time investment required for the quantitative processing of such samples, this high spatial variability in tardigrade diversity and abundance may make them unsuitable for rigorous quantitative testing of ecological hypotheses.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 45
Auto-Montage Imaging for Tardigrades
1 2 William R. MILLER and Zephyr H. JOHNSON
1Department of Biology, Chestnut Hill College, Philadelphia, PA, U.S.A.
2Biodiversity Division, Academy of Natural Sciences, Philadelphia, PA, U.S.A.
Abstract. Advances in computer-aided microscopy are going in many directions today and many of us have discovered digital imaging to be very useful for references, measurements, and comparisons. Because tardigrades are three-dimensional we understand the depth of field limitations imposed by the physics of magnified light. When using a microscope we overcome the issue by continually focusing. When we take a picture or image we are limited to the one
layer in focus and the out of focus material around our area of interest may render an image
useless. The Montage concept seems to show real some promise for tardigrades. For transparent
and three-dimensional specimens Montage imaging appears to be able to produce a clearer image
with more detail in focus than conventional or digital photography. The idea is to take a series of
digital images as layers while focusing down through the specimen. In practice, the operator
identifys the top and bottom of the specimen, selects the number of layers to be imaged, and tells
the computer to take the images. The computer controls a stepper motor that moves the focus a
fraction of a millimeter, takes an image of the layer, and moves again. Each layer image is
stored as digital data and can be viewed individually or assembled by the computer according to
an algorithm into a montage image. The montaging process analyzes each layer and erases any
part of the image that is not within the depth of field range and sharply focused. Then the
montage image is assembled using only that part of each layer that is in focus. The result is a
very three-dimensiona, in focus image that should be most useful in tardigrade taxonomic work.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 46
Tardigrades of the Sub-Antarctic: 5000 year old eggs from Marion Island
1 2 William R. MILLER and Harold F. HEATWOLE
1Department of Biology, Chestnut Hill College, Philadelphia, PA, U.S.A.
2NorthCarolina State Universtiy, Realigh, North Carolina, U.S.A.
Abstract. During the examination of pollen analytical samples taken from two peat bogs on Sub-
Antarctic Marion Island, tardigrade eggs were recognized. Samples from two sites at Albatross
Lakes and Kildalkey have yielded over 500 eggs of four different species of tardigrades. The samples were radiocarbon dated in the original research and date as far back as 7300 years before present. At the 5000-year level, tardigrade eggs appear in the samples. Cores of peat bogs represent a record of the history of vegetation and animal change or stability over time. Using the assumptions developed by pollen researchers that the debris that settles onto the bog is in proportion to its content in the atmosphere, which is in proportion to its concentration at its source. We suggest a similar model for tardigrade populations.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 47
Tardigrade Distribution in a Medium-sized City of Central Argentina
1 1 1 1 María C. MOLY de PELUFFO , Julio R. PELUFFO , Alejandra M. ROCHA and Irene L. DOMA .
1Facultad de Ciencias Exactas y Naturales, Universidad Nacional de La Pampa, Uruguay 151, 6300 Santa Rosa, La Pampa, Argentina.
Abstract. The distribution and abundance of tardigrades in the city of General Pico (35º40' LS;
63º44' LO) – Argentina – are analyzed from samples taken during autumn and spring 2001.
Sampling sites included paved urban locations with different traffic intensities, non-paved
periurban places with abundant suspended dust, and places with peculiar conditions such as the
city industrial area and the bus station. Trees of the same species were selected in each area and
from each of them nine circular samples of moss and/or lichen were taken with an 11 millimeter diameter sampling tool. The diversity, density and relative abundance of tardigrades was recorded. Species richness varied from 1 to 4. The species recorded were: Echiniscus rufoviridis,
Milnesium tardigradum, Ramazzottius oberhaeuseri, Macrobiotus areolatus and an undescribed species of the family Macrobiotidae. R. oberhaeuseri and M. tardigradum were the most frequent species. R. oberhaeuseri dominates in periurban areas with high suspension dust and very exposed to the sun. This agrees with the known higher resistance to drought of R. oberhaeuseri. M. tardigradum dominates on paved streets with intense vehicle traffic. M. areolatus and Macrobiotidae sp. are uncommon, but wherever present they are the dominant species. E. rufoviridis appears in only a few samples and always very few specimens. In a site placed near a lead polluting operation (battery factory), only two specimens of tardigrades were
found. Results support the hypothesis of a relationship between the air quality and the
lichenophile meiofauna.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 48
Tardigrade Community Composition in four forest types in the El Diviso Reserve, (Santander, Colombia)
1 1 Eliana X. NARVAEZ and Javier JEREZ - JAIMES.
1Department of Biology, University of Puerto Rico , Mayagüez ,Puerto Rico, U.S.A..
Abstract. A survey of the community composition and distribution of bryophilous tardigrades in
the “El Diviso Reserve,” Santander (Colombia), was conducted during the time period from
March 1999 through February 2000. Four forest types were chosen, old pine forest (Pinus patula) with secondary forest at 1800 m, cypress forest (Cupressus lusitanica) at 1850 m, pine forest at 1970 m and primary Sub-Andean forest at 2100 m. In each forest a plot or transect of
200 m2 was used and all trees present were sampled. Tardigrades were extracted from the
samples, mounted individually in Hoyer's medium, and identified to species using phase microscopy. Variation with altitude and in composition were determined. In El Diviso Reserve tardigrades were present belonging to 2 classes, 8 genera, and 15 species
(Calohypsibius verrucosus, Hypsibius arcticus, H. dujardini, Isohypsibius prosostomus,
Itaquascon bartosi, Pseudechiniscus novaezeelandiae, Macrobiotus areolatus, M. harmsworthi,
M. hufelandi, M. islandicus, M. cf. occidentalis, M. richtersi, Minibiotus intermedius, and
Milnesium tardigradum). Differences in tardigrade community composition and altitudinal variation were found. The cypress forest had the highest richness and diversity values. The cypress is an introduced species that turned out to be a good phorophyte for mosses and tardigrades.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 49
Long-term anhydrobiotic survival of lichen-dwelling tardigrades
Lorena REBECCHI, Roberto GUIDETTI, Simona BORSARI, Tiziana ALTIERO, and Roberto BERTOLANI
Department of Animal Biology, University of Modena and Reggio Emilia, Moden, Italy.
Abstract. It is not rare to find in references that anhydrobiotic tardigrades can survive for more than a century. However, a closer look at the empirical evidence provides very little support that tardigrades are capable to survive dried for such a long time, suggesting that 7-8 years may represent the limit of cryptobiotic survival (Jönsson and Bertolani: J. Zool. London 2001, 255: 121-123; Guidetti and Jönsson: J. Zool. London 2002, 181-187). In addition, the available data derived from limited experiences in which statistical analysis was not applied to verify the recovery rates. In order to fill this gap, we carried out a study to evaluate the long-term survival of naturally dried tardigrades. A large fragment of dry lichen (Xanthoria parietina) was collected in field after two days from rain in 1999. The dry lichen was stored inside a paper bag in laboratory at room conditions with registered humidity and temperature. Four weighed replicates of lichen were rehydrated after various lengths of storage, all tardigrades extracted and the survivors enumerated. Five species of tardigrades were found, but two of them only occasionally. Ramazzottius oberhaeuseri and Echiniscus spp. were sufficiently represented for statistical analysis. At the beginning of the experiment a percentage of R. oberhaeuseri (8.9%) and of Echiniscus spp. (28.3%) did not survive. A significant decrease in recovery of R. oberhaeuseri was observed after 86 days. Echiniscus spp. survived up to 1082 days, while R. oberhaeuseri still has 21.7% of survival after 1192 days. In addition, in R. oberhaeuseri, significant intraspecific differences in survival rate were found in relationship to the animal age, moulting and female gonad stage. A recovery after four years of anhydrobiosis should be considered a long-term survival, important from an ecological and evolutionary point of view. A wider knowledge of cellular and molecular mechanisms allowing this longevity may find biotechnological perspectives.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 50
Resting eggs in tardigrades
Lorena REBECCHI, Giorgia RINALDI, and Tiziana ALTIERO
Department of Animal Biology, University of Modena and Reggio Emilia, Modena, Italy
Abstract. Dormancy includes both diapause and quiescence. These phenomena are respectively under endogenous and exogenous control. Both dormancy forms occur in tardigrades. Nevertheless, only some species of tardigrades are able to carry out both quiescence (cryptobiosis) and diapause (encystment). Little is known on egg dormancy in tardigrades, apart their possibility to survive dehydrated. Current literature rarely refers on the production of two different kinds of eggs in the same species. In particular, thin-shelled and thick-shelled eggs were identified only in few cases. According to some authors, thick-shelled eggs are produced when environmental conditions are unfavourable. Our analysis of life history traits of a reared strain of Macrobiotus richtersi evidenced that hatching phenology is spread in about 90 days. As a consequence, we carried out a research on the eventual presence of resting eggs in M. richtersi, using an apomictic triploid cytotype. We have utilized the first oviposition in lab of females sampled in nature in spring and in the fall, and eggs from several generations of different clones reared in lab. All the laid eggs were maintained in water up to the eventual hatching. About 90% of hydrated eggs hatched, with a time of development ranging from a minimum of 30 days to a maximum of 62 days. The eggs unhatched after 90 days have been observed at LM, dried, maintained dried for 21 days and then rehydrated. Seventy-three (9.4%) eggs did not hatch and were then dehydrated. Nine of them, most of which with a completely formed animal inside, hatched after rehydration. Therefore, in this species subitaneous and resting eggs are present. The last ones need a cue to hatch, suggesting that another form of diapause may be possible in tardigrades. This represents a further strategy to colonize and inhabit unpredictable environments.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 51
Tardigrades of North America: Southeastern Pennsylvania, U.S.A.
1 1 Diana SUCHARSKI and William R. MILLER
1Department of Biology, Chestnut Hill College, Philadelphia, PA, U.S.A.
Abstract. A general survey of tardigrade habitats was conducted through out southeastern
Pennsylvania during the spring and summer of 2003. The collections included moss, lichen, soil, litter, and water at each site if available through out the southern part of the Delaware River watershed. Each location was recorded with global positioning systems for Geographical mapping. The substrate, habitat, and exposure was recorded for each sample. Tardigrades, rotifers, and nematodes were counted. Tardigrades were extracted by soaking, mounted in
Hoyer’s medium and identified to species. The collection was analyzed for patterns and relationships between and among species of tardigrades, between the other animals, their habitats, and substrates. Present in Southeastern Pennsylvania were tardigrades of 2 classes, 8 genera, and 14 species. Significant patterns of association were detected between species of animals and specific habitats. The patterns have been compared spatially with human demographic patterns.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 52
Oogenesis of Milnesium tardigradum
Atsushi C. SUZUKI
Department of Biology, University of Keio, Hiyoshi,Yokohama, Japan
Abstract. A parthenogenetic strain of Milnesium tardigradum has been maintained since autumn 2000. These animals were fed on a monogonont rotifer and grew into mature adults at the 3rd-instar stage. The first period of egg laying accompanied the third moult. The egg-laying
/moulting intervals of adult animals were around 6-10 days. The life history of M. tardigradum under the rearing environment included up to seven periods of moult or five times of egg laying.
The number of eggs in a clutch varied according to the nutritional condition of the mother and
ranged from 1-12 eggs/clutch. It is an interesting problem how the number of eggs is decided. In
this study, the internal structure of ovary was morphologically investigated to elucidate the
relationship of oocytes and other ovarian cells. Specimens from immature larvae and adults with
various stages of ovaries were fixed in glutaraldehyde and embedded in Epon. Semithin and
ultrathin sections of these specimens were observed by light microscopy and transmission
electron microscopy, respectively, and fine structures of the ovarian cells were described.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 53
Establishment of a Culture System for and Lifecycle Dynamics of the Tardigrade Hypsibius dujardini.
Fran THOMAS, Jennifer DAUB, Habib MAROON, Aziz ABOOBAKER, and Mark BLAXTER
Institute of Cell, Animal and Population Biology, Ashworth Laboratories, University of Edinburgh, Kings Buildings, Edinburgh, EH9 3JT, UK.
Abstract. We have initiated a programme of evolutionary developmental biology research on a cultured tardigrade, Hypsibius dujardini. Tardigrades are an attractive organism for comparative work because of their basal position in the pan-Arthropoda, and the observation that they share some morphological and developmental characteristics with other Ecdysozoa such as the nematodes. Previously, developmental and other research on tardigrades has been hampered by an inability to grow them in culture. We have established conditions for culture of a small, fresh water, herbivorous tardigrade using a defined, clonal food source (Chlamydomonas reinhardtii) and are able to rear tens of thousands of tardigrades. Importantly, we are also able to rear single tardigrades in multiwell plates. The culture system is based on seeding large volumes of fresh water with ample C. reinhardtii grown in Bold's medium. Aeration and illumination at 20 degC is sufficient to maintain cultures for several weeks. H. dujardini appears to be matriclonal, as we are able to rear isolated eggs or juveniles in isolation to fecund adulthood. Development time in the egg is about 4 days at 20 degC, and the juvenile undergoes several larval moults before first laying eggs at 6 days old. The animals continue to moult as adults, and lay additional clutches of eggs every 2 to 4 days thereafter, for at least 4 weeks. The eggs are laid within the shed cuticle, allowing us to easily follow a clutch of siblings that were laid and initiated development near- synchronously. In our culture system, mature adults can lay 15 or more at each moult. This is probably due to optimal nutritional conditions. We are investigating the developmental timings of eggs from different-sized clutches, and the ability of this species to undergo cryptobiosis (either desiccation or freezing).
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 54
Tardigrades of The Faroe Islands
Birna V. TRYGVADÓTTIR
Zoological Museum, Invertebrate Department, Universitetsparken 15, DK-2100 Copenhagen Ø, Denmark, e-mail: [email protected]
Abstract. The tardigrade fauna of the Faroe Islands has never been subject to thorough
investigations, and there are only a few records available from the literature. Tuxen made a
synopsis of the tardigrade fauna in 1941 based on scarce material, and his species list include 18
species. This paper presents some preliminary results from an ongoing investigation of the
freshwater and terrestrial tardigrade fauna of the Faroe Islands. Samples were collected in 2001-
2003 at several locations, habitats and altitudes on the Isles. So far the species list of hetero- and
eutardigrades contains 28 species, including a new genus of Eohypsibiidae from high mountain
moss-cushions. The collections also include the first record of the genus Amphibolus from the
Faroe Islands.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 55
Tardigrades Density and Diversity in four life zones of Costa Rica.
Jonnathan HERRERA-VÁSQUEZ
Museo de Zoología, Universidad de Costa Rica. E-mail: [email protected]
Abstract: The objective of this study is determine the diversity, average density and association between species of tardigrades of folious lichens in four life zones of Costa Rica:
Tropical moist forest (T - mf), Premontane moist forest (P - mf), Lower montane moist forest
(LM - mf) and Montane wet forest (M -wf). A total of 77 samples of folious lichens were analyzed during 2002. 14 species of tardigrades were found with 118.2 ind/ cm2 average density. The most diverse zone was the premontane moist forest (H = 2.057) and lowest low montane moist forest (H = 0.90). were observed that in three life zones the Heterotardigrada
Class is the most abundant (t = 1.9432, gl = 6, p = 0.05333). Only between Milnesium tardigradum species and Echiniscus sp2 were found significative association (p = 0.039).
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 56
Tardigrades (Phylum Tardigrada) from the western part of the Central Valley, Costa Rica with Some Ecological Annotations. 1 2 Jonnathan HERRERA -VASQUEZ & Mario Vargas VARGAS .
1. Museo de Zoología, Universidad de Costa Rica. E - mail: [email protected] 2. Facultad de Microbiología, Universidad de Costa Rica. Address: Laboratorio de Artropodología Médica (270 B), Facultad de Microbiología, Universidad de Costa Rica.
Abstract: During 2001 and 2002, tardigrades of Folious lichens from the western part of
the Central Valley of Costa Rica were collected at different altitudinal regions and fixed for
their identification. There were found four genera and seven species: Macrobiotus richtersi,
Macrobiotus harmsworthi, Macrobiotus areolatus, Isohypsibius bakonyensis Milnesium
tardigradum, Echiniscus bigranulatus and Echiniscus angolensis. The most frequent species
was M, ricthersi (31 %) and the least M. areolatus with (7.33 %) in the samples analyzed. This
is the first record of Echiniscus angolensis for Central America. There are now known 13
species of tardigrades for Costa Rica and 7 for the area of study.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 57
A Family Analysis of Tardigrade Phylogeny
1 2 1 P. Brent NICHOLS , Diane R. NELSON , James R. GAREY
1Department of Biology, University of South Florida, Tampa, FL U.S.A.
2Department of Biological Sciences, East Tennessee State University, Johnson City, Tennessee, U.S.A.
Abstract. There is no overall phylogenetic hypothesis of tardigrade evolution in the current
literature. The present study developed a morphological data set suitable for cladistic analysis at the family level. A data matrix consisting of 50 characters for 15 families of Tardigrada was analyzed by maximum parsimony. Kinorhynchs, loriciferans, and gastrotrichs were used as outgroups, and ground pattern characters for tardigrades were established from the literature.
The results agree with the currently accepted hypothesis that Eutardigrada and Heterotardigrada are each monophyletic groups. Among the eutardigrades, Eohypsibiidae was found to be a sister group to Macrobiotidae + Hypsibiidae. Necopinatidae appears to be basal among the Parachaela while Milnesiidae was the basal eutardigrade. The enigmatic Apodibius was also found to be basal among the eutardigrades but this position is inconclusive because of its lack of characters.
Among the heterotardigrades the family Oreellidae was found to be basal. Coronarctidae +
Batillipedidae were found be sister groups to Echiniscoidea + Echiniscidae. Therefore, the order
Arthrotardigrada appears to be paraphyletic and the order Echiniscoidea may be polyphyletic.
The 18S rRNA gene sequence of Batillipes was obtained and its addition to a previously
published dataset supports the monophyly of Heterotardigrada. The analysis of the
morphological data set suggests that 18S rRNA sequences from members of Oreellidae,
Renaudarctidae, and Halechiniscidae would be useful to test the paraphyletic and polyphyletic
groups that appeared in the morphological analysis.
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 58
Symposium Participants
Lois Bateman Diane Nelson Sir Wilfred Grenfell College East Tennessee State University University Drive Box 70703 ETSU Corner Brook Newfoundland and Labrador A2H Johnson City TN 37614-1710 USA 6P9 Canada 423-439-4376 709-637-6247 [email protected] [email protected]
James Garey Mark Blaxter University of South Florida ICAPB Department of Biology Ashworth Labs, King's Buildings 4202 E. Fowler Ave. SCA110 Edinburgh EH9 3JT UK Tampa FL 33620 USA +44 131 650 6760 813-974-3900 [email protected] [email protected]
Frank Romano Harry Meyer Jacksonville State University McNeese State University 700 Pelham Road N. PO 92000, Dept of Biol. Environ. Sciences Biology Department McNeese State University Jacksonville AL 36265 USA Lake Charles Louisiana 70609 USA 256-782-5038 337-475-5671 [email protected] [email protected]
Juliana Hinton Amber Hohl McNeese State University Iowa State University P.O. Box 92000 2604 Stange Rd. Apt. 7 Lake Charles LA 70609-2000 USA Ames IA 50010 U.S.A. (337)475-5651 319-470-1439 [email protected] [email protected]
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 59
Habib Maroon James Young University of Edinburgh Jacksonville State University Institute of Cell, Animal and Poulation Biology, 700 Pelham Road N. King's Buildings Biology Department West Mains Road. Jacksonville AL 36265 USA Edinburgh Scotland EH9 3JT UK 256-782-5642 44 (0) 131 650 6761 [email protected] [email protected]
Jennifer Daub Lukasz Michalczyk University of Edinburgh Jagiellonian University, Department of Ashworth laboratories Zoopsychology West Mains Road ul. Ingardena 6 Edinburgh EH9 3JT UK Krakow 30-060 POLAND +44 131 650 6761 +48126336377 ext.2461 [email protected] [email protected]
Robert DaFoe P. Brent Nichols Jacksonville State University Univ. of South Florida 700 Pelham Road N. Dept. of Biology, SCA 110 Biology Department 4202 E Fowler Ave Jacksonville AL 36265 USA Tampa FL 33620 USA 256-782-5642 813-974-8967 [email protected] [email protected]
Colleen Mitchell Jonnathan Herrera-vásquez Jacksonville State University Universidad de Costa Rica 700 Pelham Road N El Tajo casa Nº 2 Biology Dept. Esparza Puntarenas Costa Rica Jacksonville AL 36265-1602 USA 207 4193 256-782-5642 [email protected] [email protected]
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 60
Nigel Marley Jesper Guldberg Hansen School of Biological Sciences, Zoological Museum, University of Copenhagen University of Plymouth Universitets parken 15 Drake Circus DK-2100 Copenhagen East DK-2100 Denmark Plymouth Devon PL4 8AA United Kingdom 35 32 11 16 44-(0)1752-232939 [email protected] [email protected]
Frances Thomas Alexandra Avdonina ICAPB, University of Edinburgh. Vladimir State Pedagogical University Ashworth Labs, King's Buildings, Stroiteley prospect, 11 West Mains Road, Vladimir 600024 Russia Edinburgh Scotland EH9 3JT United Kingdom 10-7-0922-339784 (0131) 650 6761 [email protected] [email protected]
Peter Degma Clark Beasley Comenius University, McMurry University Faculty of Natural Sciences Department of Biology Mlynska dolina B-1 Abilene Texas 79697 U.S.A. Bratislava SK-842 15 Slovak Republic 325-793-3867 +421 2 60296492 [email protected] [email protected]
Hiroki Harada National Agricultural Research Center for Iben Heiner Tohoku Region Zoological Museum of Copenhagen Arai Universitetsparken 15 Fukushima-city Fukushima Prefecture 960-2156 Copenhagen OE 2100 Denmark Japan +45 35321039 +81-024-593-6175 [email protected] [email protected]
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 61
Birna Trygvadóttir Reinhardt Kristensen Zoological Museum of Copenhagen Zoological Museum of Copenhagen Universitetsparken 15 Universitetsparken 15 Copenhagen OE 2100 Denmark Copenhagen OE 2100 Denmark +45 35321039 +45 35321118 [email protected] [email protected]
William Miller Ruth Dewel Chestnut Hill College Appalachian State University 9601 Germantown Ave Boone NC 28608 USA Philadelphia PA 19118 USA 828 262 2682 215-248-7029 [email protected] [email protected]
Jerome Regier Roberto Guidetti Center for Biosystems Research, University of University of Modena and Reggio Emilia Maryland Biotechnology Institute Via Campi 213/d Plant Sciences Building, Rm 5140 Modena College Park MD 20742 USA Modena Italy 41100 Italy 301 405 7679 ++39 0592055555 [email protected] [email protected]
Lorena Rebecchi Roberto Bertolani University of Modena and Reggio Emilia - University of Modena and reggio Emilia - Department of Animal Biology Department of Animal Biology Via Campi 213/D Via Campi 213/D Modena 41100 Italy Modena 41100 Italy +39 0592055553 +39 0592055545 [email protected] [email protected]
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 62
Atsushi Suzuki Mary Marshall Keio University 10277 S. St. Rd. 335 Hiyoshi 4-1-1 Pekin Indiana 47165 United States of America Kohoku-ku (812)-366-3553 Yokohama Kanagawa 223-8521 Japan [email protected] 81-45-566-1329 [email protected]
Wataru Abe Daiki Horikawa Division of Biological Sciences, Graduate Department of Environmental Earth Science, School of Science, Hokkaido University Hokkaido University Kita-10, Nishi-8 Kita 10, Nishi 5, Kita-ku Kita-ku Sapporo Hokkaido 060-0810 Japan Sapporo Hokkaido 060-0810 Japan +81-11-706-2251 +81-11-706-3524 [email protected] [email protected]
Michael Collins Memorial University of Newfoundland Richard Helm Elizabeth Avenue Virginia Tech Arts-Administration Building West Campus Drive St. John's Newfoundland and Labrador A1C 5S6 Blacksburg VA 24061 USA Canada 540-213-4088 709-737-8411 [email protected] [email protected]
María Fernández Alejandra Rocha Facultad de Cs. Exactas y Naturales- UNLPam Facultad de Cs. Exactas y Naturales - UNLPam Uruguay 151 Uruguay 151 Santa Rosa La Pampa 6300 Argentina Santa Rosa La Pampa 6300 Argentina 54 - 2954 - 42 5166 54 2954 42 5166 [email protected] [email protected]
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 63
Karsten Klage Jette Eibye-Jacobsen Virginia Tech Zoological Museum, Copenhagen Biochemistry Universitetsparken 15 205 Engel Hall 2100 Copenhagen Ø Blacksburg VA 24061 USA Copenhagen Denmark Denmark 540-231-8435 +45 35321081 [email protected] [email protected]
Tiziana Altiero Matthew Boeckner University of Modena and Reggio Emilia Memorial University of Newfoundland Via Campi, 213/D Department of Biology Modena I-41100 Italy St. John's Newfoundland A1B3X9 Canada +390592055554 (709)737-8411 [email protected] [email protected]
Maggie Ray Clayton Marshall North Carolina State University Eastern High School 4277 The Oaks Drive 10277 S. St. Rd. 335 Raleigh NC 27606 USA Pekin IN 47165 United States of America 919-233-8750 (812)366-3553 [email protected] [email protected]
Paul Bartels Gallo D'Addabbo Maria Warren Wilson College Dipartimento Zoologia Università Bari WWC 6032 Via Orabona ,4 Bari Italy PO Box 9000 Bari 70126 ITaly Asheville NC 28815 USA +39 0805443345 828-771-3781 [email protected] [email protected]
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 64
Eliana Narvaez-Parra Javier Jerez-Jaimes University of Puerto Rico University of Puerto Rico Calle Mendez Vigo 165-0 apt 1007 Calle Mendez Vigo 165-0 apt 1007 Mayaguez Puerto Rico 00681-3256 Puerto Rico Mayaguez Puerto Rico 00681-3256 Puerto Rico 787-8324040 ext 2269 787-832-4040 ext 2269 [email protected] [email protected]
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 65
NOTES
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 66
NOTES
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 67
NOTES
St. Pete Beach, Florida USA 27 July – 1 August, 2003 9th International Symposium on Tardigrada 68
NOTES
St. Pete Beach, Florida USA 27 July – 1 August, 2003