~001ogicalJournalofthe Linnean So&& (1996), 116: 185-204. With 5 figures
Tardi'ade
biology. Edited by S. J. McInnes and B. B. Norman
@
Tardigrada of the Arctic tundra with
descriptions of two new species
w. MAUCCI
Via G.Mameli 9, 37100 V'erona, ItaQ"
Findings are reported from samples collected in tundra in Greenland, Iceland and the Svalbard Islands.
In total 58, tardigrade species are reported, only 12 of which are common to all three areas. Forty-three
species are reported from Greenland (1 1 new to the country and two new to science); 33 species are
reported from Iceland (14 new to the country); 21 species are reported from the Svalbard Islands (4new
to the archipelago). A description of Hypsibius pachyunguis sp. nov and Fujiscon triodon sp.
nov. (Eutardigrada, Hypsibiidae) is given.
01996 The Linnean Society of London
ADDITIONAL KEY WORDS: -Eutardigrada - Hypsibiidae - Hypsibirrs - Fujiicon,
CONTENTS
Introduction . . . . . . .
Material and Methods . . . .
Comments on the species found .
Conclusions . . . . . . .
References . . . . . . . .
. . .
. . .
. . .
. . .
. . .
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
185
186
187
202
203
INTRODUCTION
The arctic tundra, the desert that embraces the Arctic and sub-Arctic mainland
and islands beyond the area of the last dwarf trees, is a cold and generally dry
environment, with damp and relatively warm but very short summers. This type of
climate, in concert with the abundance of moss and lichen vegetation on the ground
and rocks exposed to the sun, is particularly favourable for the presence of large
Tardigrada populations, often with a great variety of species, some of which may be
endemic. Moreover, the continuity of the vegetation favours the spread of individual
species across large surfaces, instead of the 'insular' distribution of Tardigrada typical
of other environments.
Following the interesting results obtained from a great number of samples
collected throughout a large portion of the Scandinavian peninsula (Durante Pasa &
Maucci, 1975, 1979), where hundreds of specimens belonging to 60 different species
*Reprint requests to S. J. McInnes, BAS, High Cross, Madingley Road, Cambridge CB3 OET
0024-4082/96/010185
+ 19 $18.00/0
185
01996 The Linnean Society of London
186
w.MAUCCI
were found, it was decided that research should be extended to Greenland, where
two sampling expeditions took place in 1987 and 1989. Terrestrial Tardigrada from
Greenland had been previously known from two groups of locations: six on the
north-eastern coast between Bronlund Fjord, 82' 12' latitude N, and Scorebysund,
70'29' lat. N (Richters, 191la; Marcus, 1936; Petersen, 1951; Skmkria & Elin, 1989;
SkmCria, 1990); eleven along the south-western coast, between Blue West, in the
Julianhaab District, 6 1'20' lat. N, and Holsteinsborg, 65"50' lat. N (Petersen, 1951).
Other species from thermal springs were collected on Disko Island (70" lat. N)
(Kristensen, 1982; Grongaard, Kristensen & Petersen 1990).
The objective of these expeditions was to extend sampling to other areas outside
the above-mentioned locations. In 1987, samples from Southern Greenland, near
Narssaq and around the Narssarssuaq airfield were collected. These samples
contained 357 specimens of Tardigada and their eggs, belonging to 30 species, three
of which were new to science, while eight had never before been reported from
Greenland. The three new species are described in Maucci (1988).
The very promising results from this expedition led to a new sampling campaign
on the western coast, with the aim of collecting material farther north of
Holsteinborg, where Petersen's sampling had stopped. Six new locations were
visited, from Holsteinborg (65"50' lat. N) to Uppernavik (71'46' lat. N). 448
specimens in all were collected, belonging to 38 species, two of which were new to
science and a further nine new to Greenland.
A systematic survey of Tardigrada from Iceland is presented in Morgan (1980). In
1990, 29 different locations in Iceland were sampled resulting in a total of 214
mounted specimens belonging to 31 species, 14 of them new to Iceland.
After Dastych's (1 985) exhaustive work on Tardigrada from West Spitzberger, no
major discovery could be expected from this area. However, in June-July 1992 on
a cruise to the Svalbard Islands, samples from nine different locations were collected,
only two of which had been mentioned by Dastych. About 200 specimens belonging
to 19 species were found.
MATERJALS AND METHODS
This study reports the findings of the above-mentioned expeditions. The sampling
locations are as follows:
Greenland, 1987 expedition
Narssaq, in the ViUage and its immediate surroundings; Kvanefjeld; landings of
Tulleruanat and Naajat, on the banks of the Brede Fjord.
Narssarssuaq, near the afield; along the road from the airfield to the Kiagtut
glacier; Sermiat, on the front of the 'inlandsis'; Qassiarssuaq, on the Eriks Fjord;
between Itilleq and Igaliko, on the isthmus between the Eriks Fjord and the Einars
Fjord.
Greenland, 1989 expedition
Holsteinborg (Sisimiut): in the village and in the tundra of a glacial valley about
10 kms inland; Egedesminde (Aasiatt); Christianshaab (Qasigiannguit);Jakobshavn
(Ilulissat):in the village and surroundings and at Sernemiut, on the ice fjord; Umanak
ARCTIC TUNDRA TARDIGRADES
187
(Ummanaq): in the village and surroundings and at the foot of the “heart-shaped
mountain”; Uppernavik.
Iceland, 1990 expedition
Thingvellir (National Park NE of Reykjavik); Skalholt; Hellisheidi (near Hella, at
a short distance from the southern Coast); Seljalandsfoss (near the West Coast);
Skogafoss (near Skogar); Skogar; Myrdalssandur (on the coast East of Vik);
Skaftafellsysla; Eldhraun; Svinafell (near one of the tongues of the Vatnajokull
glacier); Stokkness (SE coast, radar station); Djupivogur; Hvalness (South of
Djupivogur); Bernfjordur (eastern coast, on the fjord banks); Seydisa (South of
Egilstadir, near the Seydisfjordur); Dattifoss; Asbyrgi; Godafoss (waterfall East of
Akureyri), Laxa (on the river, near the Myvatn lake); Raykiahlid (on the NE shore of
the Myvatn lake); Dimmuborgir (lavic area East of the Myvatn lake); Dalvik (near the
northern coast); Lagheidi (northern coast, between Dalvik and Hofsos); Blondugd
(inland);Audkuluhaidi (inland);Hveravellir (hot springs between the Langjokull and
Hofsjokull glaciers); Kiolur (inland, altitude: 660m); Hvitarvatn (at the foot of the
Langjokull glacier, by the lake); Selfoss (above the village of Overghamrar).
Svalbard islands, 1992 exbedition
West Spitzbergen: Isfjorden: Longyearbin and Barentsburg
Kongsfjiorden: Ny hesund
Hornsund: SkrAl Pynton
Van Keulenijorden: Ingebritsen Bukta (Whlte Whale Bay) and
Calipso Bay
Van Mijenfjorden: VArsol Bukta
Edgeoya Island Bjmnbukta
Russebukta.
At each location in Greenland from 10 to 30 samples of moss and lichen were
collected from the ground or from rocks exposed to the sun. In Iceland, from 4/5 to
about 15 samples were collected at each location (moss and lichen from the ground
or from rocks, lava and roofs), all in the sun. At each of the Svalbard locations about
20 samples from the ground or from rocks were collected, all of them in the sun.
Nearly all the above mentioned samples contained Tardigrada.
The samples were wetted for some hours and then repeatedly squeezed in a Petri
dish. The water and debris were examined under a stereoscopic 40 X microscope,
and the specimens and eggs were extracted by means of a capillary pipet. Following
live observation, most specimens were permanently mounted in polyvinyl-lactophenol and submitted to phase contrast and Nomarski interference contrast
examinations. All preparations are now included in the Maucci Collection, at the
Verona Natural History Museum.
COMMENTS ON THE SPECIES FOUND
Proechiniscus hanneae (Petersen)
This species has been reported exclusively from Greenland. Some specimens,
including an exuvia containing three eggs, were found at Jakobshavn, both near the
188
w.MAUCCI
village and at Sernermiut on the edge of the glacier. One specimen was also found
in the village of Narssaq, and a further population between Itilleq and Igaliko.
Echinlscus arctomys Ehrenberg
This species has not been sufficiently described, and is difficult to distinguish from
other similar species. Some Icelandic specimens found at Hveravellir have been
referred to E. arotomys, a species not previously known in Iceland. Although their
terminal plate is not faceted, they differ from E. wendti in that their sculpture is finer,
and the granule dotting hard to resolve even by immersion, and that their A
appendices are shorter (28-3Oo/o of the total body length). No specimens attributable
to this species were found in either Greenland or the Svalbard Islands.
Echinlscus blumi Richters
Fully developed specimens of E. blumi, complete with all the appendices, were
found at three Merent locations in Greenland (Holsteinsborg, Christianshaab and
Upernavik). In Iceland, this species was found to occur at Skalholt, Svinafell,
Reykjahlid and Dimmurborgir. The lateral B appendices are always present, even in
172 pm-long two-clawed larvae. Eschiniscus bhmi was also reported by Dastych
(1985), from two locations in West Spitzbergen, but it is not included in material
newly collected from that area. It is a widespread species, probably cosmopolitan.
Echiniscus ganulatus (Doyere)
This species is very common in Palearctic areas, and extremely rare in America
(where it has been found only in Yellowstone Park). It has not been found in
Greenland. In the Svalbard Islands, a large population was located at SkrAl Pynten
(Hornsund).
Echinlscus mediantus Marcus
Maucci (1985) proposed the adoption of this denomination for those Echinlscus
populations belonging to the blumi group in which the lateral B appendix is never
present, not even in fully developed specimens, while the C appendices appear before
D, which remain invariably shorter and are often unilateral. This opinion had
already been clearly expressed by Marcus (1936), in his comment on the species
description. A large population showing the above-mentioned features was found in
Greenland, in a sample collected near the front of the Inlandsis, in the vicinity of
Narssarssuaq. A smaller population was also located in Iceland, near Stockness; a
new record for that country.
Echiniscus merohmls Richters
The present investigation indicates that this highly widespread species appears to
be perhaps the least xerophilous of all EchinircuS species. It is one of the few species
found in samples from all three collection areas which further confirms its mainly
northern distribution (Maucci, 1980).
In Greenland, a few specimens were found at Narssaq and near Igaliko, and fairly
large populations at Holsteinsborg, Egedesminde, Jakobshavn and Upernavik.
Nearly all the specimens have the lateral B filament, and therefore belong to the socalled tfomza mecica’. In Iceland, two specimens were located at Djupivogur and a
population (without the lateral B filament) at Bondugil. As for the Svalbard Islands,
Dastych (1985)reported this species from many locations in West Spitzbergen. Large
ARCTIC TUNDRA TARDIGRADES
189
Echiniscus me-rohis populations were found at Calypso Bay (West Spitzbergen) and
on the Edge0ya Island. The tfomza suecica’ is predominant among these populations
too.
Echiniscus testudo (Doy2re)
This is a highly widespread species, markedly xerophilous and montane. It is
therefore not surprising that it proved to be quite rare in the areas considered here,
not having been reported before in either Greenland or Iceland. In Greenland two
specimens were found at Umanak and one at Christianshaab, all three of them
belonging to the tfomza tnijli.?. In Iceland a big ‘quadnijlis’, over 400 pm in length was
discovered at Reykjahlid. As for the Svalbard Islands, one quadnijiis specimen about
300 ym in length was found in a sample from StrHl Pynten (Hornsund).
Echiniscus trisetosus CuCnot
This denomination is used for those populations of the E. blumi group in which the
lateral D appendix appears before the C and always remains longer, while lateral B
never appears, or only in very few fully developed individuals. Geographically, this
species is widely but discontinuously dispersed. In Greenland, from where it had
never been reported before, a population was found in a sample collected between
Narssarssuaq and the Inlandsis, in which only one specimen, 380pm long, had
reached the E. blumi stage, that is the presence of the lateral B appendices, which are
absent in all the other individuals, ranging from 220 to 424ym in length.
Although this species had also never been reported before from Iceland, it proved
to be rather frequent in the material collected for this study. Single specimens and
populations, even large ones, were collected at Thingvellir (1 specimen), Seljalandsfoss (a fairly large population), Skogafoss (3 specimens), Myrdalssandur (1
specimen), Skaftafellisla (a small population including one two-clawed larva with
lateral D appendix and without C appendix), Hveravellir (2 specimens), Selfoss (a
large population) Skogar (1 specimen). This species was not found in the Svalbard
Islands, nor has it been reported by other authors.
Echiniscus wendti Richters
This is the most common and widespread species of the so-called ‘arctomys group’.
It is quite common in Greenland, where some E. wendti populations were found in
Narssaq (mistakenlyreported in Maucci, 1988, as E. arctomys) and near Narssarssuaq,
as well as at Holsteinsborg, Egedesminde,Jakobshavn and Upernavik. In Iceland, a
specimen was found at Seljalandfoss and one population at Thingvellir. No
specimens of E. wendti were found in the Svalbard Islands, from where it was however
reported by Marcus (1936), Weglarska (1965) and by Dastych (1985).
Testechiniscus spitzbeqens2s (Scourfield)
This is an Arctic, sub-Arctic and montane species. Although it was found in all
three areas under consideration, it was rare, and the species must be far more
common than these results seem to suggest. In Greenland just one specimen was
found near Narssassuaq, while in Iceland only two specimens were found, one at
Thingvellir and one at Selfoss. In the Svelbard Islands too, only one specimen was
found at Skra Pynten. The species was described by Scourfield (1897) from West
Spitzbergen, and subsequently mentioned also by Richters (1903, 1904b, 191 1b),
Weglarska (1965) and Dastych (1985), who reported it from several locations.
190
\v. hlAUCCI
Bryodelphax sinensis Pilato
One specimen found at Holsteinsborg is new for Greenland. No specimens were
found in Iceland or in the Svalbard Islands, although Dastych (1985) reported a few
specimens from Atomfjella.
Hypechiniscus gladiator (Murray)
In Greenland, a small population was found in the village of Narssaq. In Iceland
(no previous report) a very large population was found at Thingvellir, including also
one specimen belonging toforma bigladii and two of thefomzafissigladiispecimens. The
species has never been found in the Svalbard Islands.
Pseudechiniscus islandicus (Richters)
This fine species, described by Richters (1 904), from Iceland, has been reported
from very few other locations. Its presence in Greenland was reported by SCmkria
and Elin (1989) from Scorebysund, but it was not found in material collected for this
study. In Iceland, on the contrary, P. klandicus has proved to be one of the most
common and most abundant species. Very large populations were collected at
Thingvellir (4 samples). Skaftafellsysla, Eldhraun, Djupivogur and Lagheidi. The
biggest specimen is 616pm long, and 16 out of the 47 mounted samples exceed
500 pm. P. islandicus has never been reported from the Svalbard Islands.
PseudechinZrcus suillus (Ehrenberg)
Petersen (1951) described P. suillus yoma facettalk’ from Greenland. It is here
considered that the denomination P.facettulis would be more appropriate, as it is not
possible to determine with any certainty whether this form is subordinate to P. suillus
or to P. pseudoconijii It might be either a P. suillus with faceted cephalic and terminal
plates, or a P. pseudoconfir without lateral conical appendices. Both assumptions are
equally likely. The true P. suillus has therefore never been reported from
Greenland.
In Iceland, P. suillus is one of the few species reported by Richters (1904a) and was
found at Seljialandsfoss, Skogafoss and Hvalness. In the Svalbard Islands, the species
was reported by Dastych from two West Spitzbergen locations. At Russe Bukta, on
the Edgeerya Island, two specimens were found with faceted terminal plates, which
have been identified as P. suillus because of the finely granulated cuticle.
PseudechinZrcus uiclor (Ehrenberg)
This is a holoarctic species, with a sub-Arctic and alpine distribution, reported
from Greenland by Marcus (1 936) and Petersen (1 95 l), but not included in samples
collected for this study. In Iceland large populations were found at Audkuluheidi and
Hveravellir. This species was not found in the Svalbard Islands, from where,
however, it was reported by Richters (1 9 1 1 a), Weglarska (1965) and Dastych
(1985).
Marrobiotus areolatus Murray
This species is widely but discontinuously distributed. It is very common in some
areas, where it may even be the most frequent Tardigrada species (North-East Italy,
Portugal), and very rare or even absent in others (Turkey, Scandinavian countries).
Petersen reported it (as M . richtersi, type 2) from two locations in Greenland
ARCTIC TUNDRA TARDIGRADES
191
(Nakajanga and Brerdlund Fjord), but no specimens were found in material collected
€or this study.
Fairly large populations, though not very frequent, have been reported from West
Spitzbergen in the Svalbard Islands (Marcus, 1936; Weglarska, 1965; Dastych,
1985), where only one specimen was collected (SkrH1 Pynten, Hornsund).
Macrobiotus crenatus Maucci
This species is described in Maucci, 1988 (1991). Though very similar to M.
aerolatus, it differs from it in the absence of eyespots, the presence of large dentate
lunulae on the fourth pair of legs, and in the remarkably higher and more tapering
processes of the eggs. The specimens were collected between Narssarssuaq and the
Inlandsis.
Macrobiotus crenulatus Richters
= M. echinogenitus Richters part.
= M. dentatus Binda
One population collected at Upernavik, and mainly characterized by the presence
of markedly dentate lunulae, must be ascribed to this species. Petersen (1951) uses the
name of M. echinogenitus for specimens with both crenate and highly dentate lunulae.
Specimens of the latter, to be ascribed to M. crenulatus, were found at various
locations in Greenland: one on the north-eastern coast (Clavering 0),and 5 on the
south-western coast. This species was also found in Iceland (Svinafell and
Hveravellir) but without any eggs. In the Svalbard Islands, this species was found at
one location on West Spitzbergen (Virsol Bukta, on the Van Mijenfjorden coast),
and large populations were also found at two locations on Edgeoya Island. Weglarska
(1 965) and Dastych (1985), reported M. echinogenitus from several West Spitzbergen
locations, but, as there is no mention of the lunulae, it is impossible to determine
whether some of their specimens could perhaps be attributed to M. crenulatus, as
seems very likely.
Macrobiotus echinogenitus Richters
According to commonly accepted opinion, M. echinogenitus differs from the species
described above in that it has smooth or only slightly crenate lunulae. The specimens
collected at Egedesminde and at Christianshhab, as well as those described by
Petersen (195 I) from Br0nlund Fjord, must therefore be attributed to this species,
while the identification of the Scorebysund specimens (Marcus, 1936) remains
uncertain. This species was not found in Iceland, nor in the Svalbard Islands. As to
the possible presence of M. echinogenitus in the Svalbard, see remarks about M.
crenulatus.
Macrobiotus hamworthi Murray
This is a highly widespread species, very common in all the three areas under
consideration, probably cosmopolitan. In Greenland, populations complete with
eggs were found at Holsteinsborg, Egedesminde, Umanak, Jakobshavn, Upernavib,
Narssaq and Narssarssuaq. In Iceland, many specimens and eggs were found in
samples collected at Thingvellir, Skalholt, Hellisheidi, Dyupivogur, Seydisa, Asbyrgi,
Hveravellir. M. hamsworthi was found in West Spitzbergen (Hornsund, Van
192
\v.
hlAUCCI
Keulenfjorden, Van Mijenfjorden), and also on the E d g e ~ y aIsland. One egg,
collected at Virsol Bukta, is to be ascribed to the subspecies M. obscurus.
Macrobiotus hufelandi Schultze
This very frequently reported species has recently been subdivided into as many
as about ten different species. This is due to the assumption, firmly maintained by
some authors, that intraspecific variety is extremely low among Eutardigrada.
However, this assumption has not been sufficiently demonstrated. It is felt therefore
that the introduction o f a new denomination every time a difference is observed (be
it referred to the buccal apparatus or the claws, or the egg processes, or to biometric
values) will lead to confusion rather than clarity.
O n the basis of the current classification, in Greenland no specimens were found
that could be ascribed with any certainty to the true M. hujlandi, as redescribed by
Bertolani & Rebecchi (1993). However, this species was found at 12 locations in
Iceland (Thingvellir, Skalhot, Skogafoss, Seljalandsfoss, Eldhraun, Stokness, Seydisa,
Dettifoss, Dalvik, Lagheidi, Blondugd, Hveravellir): there were a few eggs, but always
very- few specimens. No specimen of M. hufelandi, nor of the other similar species were
found in the Svalbard Islands. Dastych (1985) reported it as fairly frequent and
abundant in West Spitzbergen: considering the date of his publication, however, it is
impossible to ascertain to which species it refers.
Macrobiotus islandicus Kichters
This species is surely typical of tundra environments, and has been frequently
reported from all the three areas under consideration. In Greenland, abundant
populations and many eggs were found at Narssaq, Narssarssuaq, Jakobshavn, and
Upernavik. In Iceland, it was reported (and described) by Richters (1904a); in this
study it was found at eight locations (Thingvellir, Seljalandsfoss, Eldhraun,
Djupivogur, Seydisa, Lagheidi, Blondugil, Kiolur).
Dasqch ( I 985) reported it as very frequent and quite abundant from many West
Spitzbergen locations. It was included in the samples from Virsol Bukta and Calypso
Bay.
ilfacrobiotus macrocalyx Bertolani
Among the different species derived from the fragmentation of M. hujlandi, M.
macroca&x is the easiest to identlftj, at least as far as its eggs are concerned. It was not
found either in Greenland or in the Svalbard Islands, whereas in Iceland two small
populations with some eggs were found at Hvalness (1 specimen and 1 egg) and
Selfoss (3 specimens and 2 eggs). A quick review of the author’s collection revealed
some specimens which could be attributed to this species from Italy (Cansiglio
plateau and Val d’Aosta), Greece, Turkey and Portugal. The two pre- Kiss fossil eggs
reported in Durante Pasa and Maucci (1972) must also be attributed to M.
mnrrocalvx.
Mauobiotus persirnilis Binda and Pilato
This species, recognizable by the egg shell, which is smooth between the processes,
has been reportcd occasionally from some European locations (Italy, Greece, Spain,
Portugal, Germany, Russia). Some findings from Turkey, North Africa and Australia
seem to suggest a much wider geographical distribution. It is a new species for
ARCTIC TUNDRA TARDIGRADES
193
Greenland, where it was found near Jakobshavn. It was not found either in Iceland
or in the Svalbard Islands.
Macrobiotus peterseni Maucci
So far, this species has been reported only from Greenland, where it is quite
abundant. Besides the two locations near Narssaq and Narssarssuaq from where it
was originally described, one egg was found also at Holsteinsborg, and large
populations with many eggs in several samples from Jakobshavn. It has not been
found in Iceland, nor in the Svalbard Islands.
Macrobiotus recens Cutnot
This species is very common in Southern France and in the Iberian peninsula, and
its presence in Greenland could therefore be surprising. It was reported from
Scorebysund (SCmtria & Elin, 1989) under the name of M. hufelandi recenr. The
presence of this species in Greenland is however confirmed by its occurrence in
samples at Christianshaab and Jakobshavn.
Macrobiotus richteni Murray
This cosmopolitan and generally abundant species does not appear to be frequent
in tundra environments. Petersen reported it from three locations in Greenland (M.
richtersi type 1). One specimen was found at Holsteinsborg and a small population
near Narssarssuaq. The species is rare also in Iceland: just one specimen was
collected at Skogafoss. It was not found in the Svalbard Islands, from where Dastych
reported it as infrequent and limited in number.
Macrobiotus spectubilk Thulin
This species, although infrequent, is however very typical of sub-Arctic (Siberia,
Scandinavia, Alaska) and montane environments (Tatra, Alps, Caucasus). It is
reported here for the first time from Greenland: just one finding (a small population
with eggs) from Jakobshavn.
Macrobiotus willurdi Pilato
This is a new species for Greenland, but it proved to be one of the most abundant
species to be found along the western coast (Holsteinsborg,Jakobshavn, Upernavik),
with extremely large populations and many eggs contained in ten different samples.
It has never been found in Iceland, but was reported by Dastych (1985)as abundant
and quite frequent from 18 locations in West Spitzbergen. A small population was
present also in one sample from the Hornsund.
Minibiotus intemnedius (Plate)
This is a highly widespread species, which shows no particular environmental
preferences. Petersen (195 1) had already reported it from Greenland (Kangamiut),
and a small number of specimens were found at three locations in this study
(Narssaq, Narssarssuaq, Holsteinsborg). From Iceland it was reported by Richters
(1904a), and small populations were present in three samples from Seljalandfoss.
Dastych (1985)reported it as infrequent and limited in number in West Spitzbergen,
where it was not found in this study.
Rkhkrszus coronfu (Richters)
This major species, limited exclusively to Palearctic environments and particularly
frequent in montane areas, appears to be present, but not abundant, in all the three
areas under consideration. It was reported only once from Greenland by Petersen
(195 1) (Brcanlund Fjord), and was not present in material collected from that country.
In Iceland, large populations were found at two locations only (Skogar and Selfoss).
In the Svalbard Islands, two populations were collected at SkrAl Pynten and VArsol
Bukta. Richters (1903; 1904),Weglarska (1965), and Dastych (1985) reported it from
several locations in West Spitzbergen.
Am$hibolus nebulosus Dastych
This evidently sub-Arctic species has so far been reported only from West
Spitzbergen (Dastych, 1983, 1985). However, the embryonate eggs found near
Hammerfest, in Norway, and mistakenly reported as Isolypsibius smreczinskii in
Durante Pasa and Maucci (1979),must also be attributed to A. nebulosus. A very large
population with many eggs was present in a sample collected in the village of
Narssaq.
Calohypsibius caelatus (Marcus)
It has been considered by nearly all authors as a ‘form’or sub-species of C. ornatus,
both because of the authority of Marcus himself, who described it as a ‘form’, and
because of Barto? (1 940) remarks on the variability of C. ornatus. The present author
has always been in doubt about the 32 variations described by BartoS: it seems
strange that Barto? may have had at his disposal so many populations of this species
which, albeit not rare, is certainly not common, as to be able to observe so many
variations, and even to draw some conclusions of an ecological nature. Furthermore,
the present author has never been able to observe even one of all these variations. All
the specimens in my collection, from different locations, consist either of typical C.
ornatus (Belgium,Norway, Val d’Aosta, Switzerland, Portugal, Greenland, Iceland) or
of C. caelatus (Sweden, Portugal, Greenland). The same applies to the Ramazzotti
collection, which contains typical C. ornutus specimens (Val Bognanco, Switzerland,
Austria, Lapland, Black Forest) and only one C. caelatus specimen (Black Forest).
Pilato, Claxton and Binda (1989) declared C. caelatus a valid species, on the basis not
only of the different dorsal processes, but also of the different length of the claws.
This opinion is here concurred with, and consequently the single specimen found
near Jakobshavn is ascribed to this species. The remarks by Pilato et al. (1989)on the
length of the claws are also confirmed in this study. Data obtained from C. ornatus (4
specimens: 1 from Greenland and 3 from Iceland) and from C. caelatus (4 specimens:
1 from Greenland and 3 from Portugal) are as follows: the mean value for claws IV
is 28.55 pts (standard deviation: 1,3324) for ornatus and 38.465 pts (standard deviation:
2.32) for caelatus. The difference is statistically significant with a 99% degree of
probability (Student’s t-test = 5.241 1 for 6 degrees of freedom).
Calohysbius ornatus (Richters)
Only one specimen of this species was found in a sample collected between
Narssarssuaq and the Inlandsis, and another near the village of Jakobshavn (in a
different sample, collected at some distance from those containing the species
described above). There are no other reports of C. ornatus from Greenland, except
one b y Marcus (1936) who gives no further details. In Iceland a fairly large
ARCTIC TUNDRA TARDIGRADES
195
population was found at Selylandsfoss. This species was not found in material from
the Svalbard Islands, from where it was however reported by Marcus (1936) and
Dastych (1985).
Hypsibius allisoni Horning, Schuster & Grigarick
A review of some material from New Zealand (kindly presented to the author by
Nelson and Schuster) led to the conclusion that this insufficiently described species
(Horning et al, 1978), very similar to H. dujardini and H. convergens, differs from the
former in that it has no septula, and from the latter because of the presence of a
cuticular bar between the claw bases on legs IV. Specimens showing these features
(two rod-shaped macroplacoids; no septula; large claws especially on the fourth pair
of legs and presence of a cuticular bar) were found in samples from Jakobshavn and
Upernavik. Besides New Zealand, this species, which is new for Greenland, has also
been reported from the Andes region in South America.
Hypsibius convergens (Urbanowicz)
This very common and probably cosmopolitan species has already been reported
from Greenland. Populations were found near Jakobshavn and Upernavik. In
Iceland, a few specimens were collected at Skogar, Dimmuborgir and Hveravellir. As
regards the Svalbard Islands, the species was reported from West Spitzbergen by
Weglarska (1965), and (as quite frequent and abundant) by Dastych (1985).
Hypsibius dujardini (Doykre)
This is a widespread, generally aquatic species; not infrequently it is also found in
mosses. In Greenland, a large number of specimens (including also cysts and one
ovigerous exuvia) were found near Jakovshavn and at Qassiarssuaq, on the Eriks
Fjord. No specimens were found in Iceland, and only one in the Svalbard Islands,
from where H. dujurdini was reported by Richters (1904b), Murray (1907), Weglarska
(1965) and Dastych (1985). The specimen was found at Ny AIesund.
Hypsibius microps Thulin
Some populations were found in Greenland at Narssarssuaq, Holsteinsborg and
Jakobshavn. No specimen was found in Iceland, while just two specimens were
present in samples from Skril Pynten: no other report exists from the Svalbard
Islands.
Hypsibius pachyunguis sp. nov. (Fks 1-3)
Hologpe. Length: 424 pm. Colourless, smooth cuticle, eyespots absent. The buccal
tube is short (33.2pm) and narrow (2.4). The bulb contains two rod-shaped
macroplacoids, long and thin (8 and 6 pm respectively), the first with a light incision
at mid-length. Septula and microplacoids absent.
The legs are short and stocky, with big and very heavy claws. The external claws
are 20 pm long on the I pair of legs, and 25 pm long on the IV.Internal claws are
significantly smaller, but still very bulky. The basal branch of the external claws is
cylindrical in shape (5pm both in length and width), and has an expanded base; that
of the internal claws is also large, but with a somewhat smaller base. The secondary
branch is very arcuate and strong. The main branch (the connection of which to the
claw is flexible, but not in a very apparent manner) is also large and strong, with
strong accessory spines. The claws on the fourth pair of legs are longer, and their
accessory spines are more evident. An S-shaped cuticular ridge between the bases of
the claws is present on all legs.
Puru&pes. Five paratypes were found, varying in length from 340 to 400pm. Their
features correspond to those of the holotype.
The holotype (recorded as no C.T. 14375) and four paratypes (all included in the
Maucci Collection, at the Museo Civic0 di Storia Naturale, Verona) were found
around Umanak; one additional paratype was found at Jakobshavn.
Figures 1- 3. H y p s i b i w p a c h p g u i s sp. nov. Fig. 1. Buccal apparatus. Fig. 2. Claws I. Fig. 3. Claws
11.
ARCTIC TUNDRA TARDIGRADES
197
H. pachyunguis belongs to the group of species which can be defined as the ‘dujardini
g~oup’,including the colourless Hypsibius, with smooth cuticle and two rode-shaped
macroplacoids. However, it differs clearly from all the other species belonging to this
group (allisoni, antarcticus, articus, convergens, dujardini, pedrottit) in the shape of the claws,
and especially in the very strong basal branch.
Hypsibius palldus Thulin
In Greenland H. pallidus populations were found at Holsteinsborg and near
Jakobshavn. In Iceland, one specimen only was found at Lagheidi, and a large
population near the Laxa river. The species is new for the island. The species was
found in the Svalbard Islands. Weglarska and Dastych reported it from West
Spitzbergen as infrequent and limited in number.
Hypsibius scabropygus CuCnot
This species was not found, nor has it been reported by other authors, in either
Greenland or the Svalbard Islands. One specimen was found in a sample collected
near Hvitarvatn in Iceland.
Ramazzottius cataphractus Maucci
This rare species had so far been reported only from two locations: Grossglockner
(Austria, altitude: 2364m, loc. type. Maucci, 1974), and Bunsow Land (West
Spitzbergen, Dastych, 1985).A very large population was found on the rocks at the
Naajat landing, on the Bredefjord, near Narssaq. As in the case of Dastych’s
material, in these specimens the dorsal sculpture is less apparent than in the type
material.
Rarnazzottius oberhausem’ (Doy2re)
The present author has always thought that the decision to transfer R. oberhauseri,
the type-species of genus Hypsibius, to another genus was too hasty and not
sufficientlyjustified, as well as in contrast with both the letter and the spirit of the
International Code of Zoological Nomenclature. In its opinion no1551 (September
1989), the International Commission on Zoological Nomenclature decided,
following a majority vote, that dujardini was to be considered as the new type-species
of the Hypsibius genus, which - it was remarked - “seemed intended to conserve
the authors’ new genus Ramazzottius Binda and Pilato, 1986”. Though this author
remains unconvinced he would not oppose the Commission’s authority.
R. oberhauseri was reported by Petersen (1951) from Nakajanga and Brranlund
Fjord, and by StmCria and Elin (1989) from Scorebysund. A large population was
found in the immediate vicinity of Narssarssuaq airport, and a few specimens in the
village of Narssaq and at Chrieanshaab, Jakobshavn, Umanak and Upernavik. In
Iceland, a small number of specimens were collected at Reykjahlid and Audkuluheidi. This species is not included in material from the Svalbard Islands, but was
reported as quite common in West Spitzbergen by Marcus (1936),Weglarska (1965)
and Dastych (1985).
Hebesuncus conjungms (Thulin)
In Greenland, only one specimen of this species was found near Upernavik. In
Iceland it was found at two different locations (Thingvellir and Berufjordur), but was
M’. hlAUCCI
198
absent from material from the Svalbard Islands, though Dastych (1985) reported it
from one location in West Spitzbergen (Atomfjella).
Diphascon chilmense (Plate)
There exists some taxonomic confusion regarding some Diphascon species, due to
the fact that many authors have misinterpreted the features of D. alpinurn, Murray
1906. Pilato and Binda (1977), rightly emphasized that the original description ofthis
species does not mention either microplacoids or septula, nor are they present in the
original drawing, These authors therefore believe that the specimens with
alpinurn auct.) should be identified as D. pingue
microplacoids and septula (0.
(Marcus).
In fact, the latter species has significantly longer and clearly rod-shaped
macroplacoids (Marcus, 1936, compares them explicitly to those of D. prorsirostre),
growing in size from the first to the third. Consequently, the specimens with
macroplacoids shaped like short oval rods or elongated granules more or less of the
same length or only slightly growing in size should, in my opinion, be ascribed to D.
chilmense.
The specimens Petersen (1 95 1) described as D. alpinurn have both microplacoids
arid septula, and cannot, therefore, belong to this species. It seems that the specimens
described as Type A should in fact be identified as D. chilenense, while Type E is most
likely D. pingue.
Both species were present in samples from Greenland: D. chilenense in several
samples from the village of Narssaq and its surroundings, the Narssarssuaq airport
area, and the area near Jakobshavn. In Iceland, D. chilenense was collected at
Thingvellir, Asbyrgi and Lagheidi (with 40 and 50 p t long macroplacoids). This
species was not found in the Svalbard Islands. Dastych reported D. pingue (as
interpreted by Pilato and Binda, 1977) from a small number of locations in West
Spitzbergen. At least the specimen he photographed (Dastych, 1985, tab. XIV a),
however should be ascribed to D. chihense.
Dzphascon pingue (Marcus)
One population collected near Holsteinsborg and one specimen from near
Upernavik can undoubtedly be identitied as D. pingue. In Iceland, a small number of
specimens were found at Thingvellir and Magheidi. In the Svalbard Islands, a single
specimen was collected at V%rsolBukta.
Regarding the difference between D.p i w e and D. chihense, the data derived from
the author’s material (macroplacoids row in pt) are as follows:
chihense
pingue
n=7
n=7
mean 41.87; SD 2.16
(Greenland 3; Iceland 3; Svalbard 1) mean 54.1 1; SD 3.01
Student’s t-test = 6.1808 (for 12 degrees of freedom).
Diphascon prorsirostre Thulin
‘This species is very similar to D. p i q u e in terms of general appearance and
macroplacoid length, but differs from it significantly in the absence of microplacoids
and septula. The third macroplacoid often shows a caudal thickening similar to that
of D.chvutum (though this latter species has a cuticular granulation at the caudal end
which is never found in prorsirostre). Much less significant are the differences between
ARCTIC TUNDRA TARDIGRADES
199
D. prorsirostre and D. ardutjons, the specific distinction of which seems far from
proven.
This species is new for Greenland, where a single specimen was found near
Narssarssuaq airport, and several specimens in four dif€erent samples from near
Jakobshavn. It is a new species also for Iceland, where it was found at Djupivogur
and Lagheidi. It has never been reported from the Svalbard Islands. Dastych,
however, reported D. ardutjons as a rare species found at four locations in West
Spitzbergen.
Diphascon recamieri Richters
This species was not found in either Greenland (although Petersen (1951) reported
it from Itivdlinguaq), or Iceland. In the Svalbard Islands, large populations of this
species were collected at Ny kesund and Skril Pynten.
Diphascon tenue Thulin
This fairly rare species has never been reported from either Greenland or Iceland.
It is however present in the Svalbard Islands, where Dastych (1985) reported it as
infrequent and limited in number from some locations in West Spitzbergen. Material
collected for this study includes a specimen from Viirsol Bukta.
Fujscon belgicae (Richters)
( = F. diphasconiellum Ito. 1991)
The new species described by Masamichi Ito (1 99 1) is, as the author himself says,
very similar to D. bel&cm, from which it is reported to differ not only in the structure
of the claws (on the basis of which the Fzjiscon genus was established) but also in the
presence of cuticular bars near the bases of all the claws. Such bars are reported, or
shown in drawings and photographs of D. begcue, by several authors (Bernard, 1977;
Dastych, 1980 and 1988; Maucci, 1986; Weglarska, 1968; Ramazzotti & Maucci,
1983). In the specimens included in the author’s collection (from Austria, Norway,
Iceland, Svalbard Islands, as well as large populations from the Tuscan Apennines
and from Greenland), the bars on the legs are always present, and the claws are
invariably subdivided into basal, secondary and main branches by internal septa.
This separation into three branches is also evident in the photographs published by
Dastych in 1980 and 1988. Indeed, it would be too much to believe that so many
different authors actually never saw the true D.bel@cae, but always only Ito’s new
species. It seems therefore reasonable to propose F. diphconiellum It0 as a recent
synonym of F. belgzcae.
As far as the F z j k o n genus is concerned, it could perhaps be accepted in the light
of the high taxonomic value currently attached to the claw structure of Eutardigrada.
The attribution of this genus to the Eohypsibiidae family, however, fails to convince.
Although according to the letter of the family diagnosis (Bertolani & Kristensen,
1987), this attribution would seem justified (excepting the 180’ rotation capability of
the internal claw), the overall features of F. b e k c m suggest a different conclusion. The
buccal-pharyngeal apparatus is very clearly of a Diphascon type (very similar to that
of other species belonging to this genus, such as F. scoticum and F. h&mz). Regardless
of their subdivision into three branches, the claws are of the Hypsibiw type, with a
sickle-shaped basal + secondary branch system and a flexible main branch joint,
while the internal and external claws are significantly different. Consequently, the
200
\v.
hlAUCCI
hypothesis that this species may belong to the same phyletical branch as Eohpsibius
or even Amphibolus seems totally groundless.
F. be&ue was reported by Stmeria (1990), from Scorebysund. A large population
was found near Upvernavik. In Iceland a single specimen was found at Stokkness,
and a further single specimen in the Svalbard Islands at V2rsol Bukta. Both Richters
[191 1) and Dastych ( 1985) reported this species from West Spitzbergen, but it is
rare.
Fujiscon triodon sp. nov. (Figs 4, 5)
Holu&pe. Lenc#: 264 pm. Shape: slender, cylindrical, with tapering head; mouth in
a terminal position. Colourless. Smooth cuticle. Eyespots absent. The buccalpharyngeal tube is thin (2.1 pm) and has an overall length of 64.5 pm (31.5 for the
rigid buccal tube). The stylet supports (markedly curved) are inserted at 6 1.9pt. The
stylets are thin, with a large furca. The ‘drop-like’ formation is missing. The bulb is
elongated (39 X 12 pm). The apophyses are punctiform and exceedingly small.
There are three very thin macroplacoids respectively 6,5.25 and 9 pm in length. The
total row length is 22.5pm. The microplacoids are very small and thin. Septula
absent.
The claws of each leg differ greatly from one another. The external claw has a
strong common branch, protruding laterally into a small basal spur; the main branch
Figires 4 & 5 . Fujiscon trtodon sp. nov. Fig. 4. Buccal apparatus. Fig. 5. Claws 111.
ARCTIC TUNDRA TARDIGRADES
20 1
is long, thin and has small accessory spines; the secondary branch is strong and
moon-shaped. The claw complex is 15 pm long. The internal claw is smaller
(9.75 pm) and rather heavier, with a basally tapering common branch. All the claws
have internal septa clearly separating the branches as in Fujiscon bekcae. There is no
cuticular thickening at the claw base.
The holotype - recorded under no C.T. 14339 - was found in a moss sample
on a rock in the sun, on the edge of the Jakobshavn village.
Paragpes. Three paratypes were found in another moss sample collected near
Jakobshavn, but at a shorter distance from the ‘Ice fjord’, at Sernemiut.
Remarks. While the buccal-pharyngeal apparatus as a whole is virtually identical to
that of Diphascon scoticum, the claws differ greatly from those of the latter species, both
because they are subdivided into three branches as is typical of the Fujkcon genus, and
in terms of their general shape. In addition, there is no cuticular thickening on the
legs.
The main difference with respect to Fujiscon be&& is the presence of three
macroplacoids (they are three distinct placoids, and not the result of a fragmentation
of the first or second placoid). Moreover, the claws are significantly different in
shape, and have no cuticular thickening.
Also in this case, while it is obvious that the species belongs to the genus Fujiscon,
the general structure of the buccal-pharyngeal apparatus and the shape of the claws
clearly indicate that the family is Hypsibiidae, and certainly not Eohypsibiidae.
Platicrista angustata (Murray)
Only one specimen was reported by Petersen (1 95 1) from Greenland (Frederikshaab). A fairly large population was found (including one exuvia with 2 1 eggs) near
Jakobshavn. It has never been found in Iceland. In the Svalbard Islands, two
specimens were found at Skril Pynten, and the species was reported from West
Spitzbergen by Murray (1907) and Dastych (1985).
Isohypsibius glaber Durante and Maucci
Only one specimen of this rare species, previously reported only from its type
locale (Javre, Sweden), near Jakobshavn. It was not found either in Iceland or in the
Svalbard Islands.
Isohypsibius lunulatus Iharos
This is one of the best described and most easily recognizable species of the socalled ‘tuberculatusgroup’ It is also one of the most widespread tardigrade species, and
has been reported from many locations in the northern hemisphere, both in Eurasia
and in America. However, some specimens were also found in the Chilean Andes.
No specimens were found in either Iceland or the Svalbard Islands, while in
Greenland two populations were found at Jakobshavn and Upernavik.
Isohypsibius prosostomus Thulin
This is a widespread and common species. In Greenland, it was found at
Holsteinsborg,Jakobshavn, Umanak, Upernavik and Narssarssuaq. In the Svalbard
Islands, large populations were found at Skril Bukta and VArsol Bukta. Dastych
(1985) reported it as infrequent, but rather abundant in West Spitzbergen.
‘02
W.XlAUCCI
Iso/ypsibius s a t t h (Richters)
This is a cosmopolitan species, frequently reported by several authors as 1.
bakonyensis. Occasional specimens were found at Narssarssuaq and Jakobshavn in
Greenland. It was not included in material from either Iceland or the Svalbard
Islands, though Dastych (1985) reported it (under the name of I. bakonyiemis) from
West Spitzbergen as a rare species.
Milnesium eurystomum hlaucci
The typical location of this very interesting species is Narssarssuaq, in Greenland.
Other specimens were also found in the Argentinian Andes (Onelli glacier) and in
Chile (Punta Arenas). It does not appear to be present either in Iceland or in the
Svalbard Islands.
,I.lilnesium tardigradurn Doykre
A cosmopolitan and very common species, found at many locations in all three
areas under consideration.
CONCLUSIONS
’The association of a considerable number of different species in the same location
is frequently found particularly in Greenland and in the Svalbard Islands.
Greenland: Narssaq (1 3 species), Holsteinsborg (13 species), Narssarssuaq (1 1
species near the airfield and 16 species on the front of the Inlandsis),Jakobshavn (22
species), Uppernavik (1 4 species).
Svalbard Islands: Skriilpynton (1 3 species), V h o l Bukta (1 1 species).
In most cases, however, the number of individuals for each species is limited. This
could be due to the extension and continuity of the moss layer, which favour a wide
scattering of the populations and many represent an obstacle to high concentrations
of individuals, more common in isolated moss cushions. This feature is much less
evident in Iceland, where isolated moss cushions are more frequent. An exception is
represented by Thingvellir where 1 1 different species were found.
Very abundant populations were however also found, for example:
Echiniscus granulatur at SkrHlpynton; Echinlscus merokmis at Russe Bukta and
C hristianshaab; fihiniscus wendti at Narssaq; Echinkcw trisetosus at Selfoss; Hypechiniscus
gladiator at Thinkpe1lir; Pseudechinircw islandicus at Thingvellir and Djupivogur;
A4acrobiotus crenulaius at Russe Bukta; Isofppsibius prosostomus at Unianak; Macrobiotu 5
idandieus at Egedesminde; Macrobiotus peterseni at Christanshaab and between Itilleq
and Igaliko; hlacrobiotus willardi at Christianshaab and between Itilleq and Igaliko;
Adoribiotusgranulatus at Narssaq; Ramauottius catuphractus at Naaiat; Amphibolus nebulosus
at Narsaq.
Although the three areas considered are very similar in terms of climate and
vegetation, their Tardigrada populations differ considerably: only 12 of the 58
species found are present in all the three areas. Of these 12 species, five are virtually
cosmopolitan: EchinzJcus merokensls, Echtniscus &stud0 (frequently xerophilous and
mainly montane), iZlacrobiotw hamworthi, Hypsibius convergens and Milnesium tardigradurn. Testechiniscus spitzbergemis, Macrobiotus klandicus and Macrobiotus crenulatus are
Arctic, sub-Arctic and montane. Richtersius coron$er and Fuj;rcon belgicae are present
exclusively in holoarctic regions, where they are quite common; Dtphascon pingue, a
ARCTIC TUNDRA TARDIGRADES
203
holoarctic species, has been reported also from the Antarctic, while Diphascon
prorsirostre, which is also holoarctic, has been reported from New Zealand and Tierra
del Fuego.
As to the other species, ProechiniscUs hannae, Macrobiotus crenatus, Macrobiotus peterseni,
Hypsibius pachyunguis and Fujiwon triodon, found in Greenland, seem to be limited
exclusively to that area, while Pseudechiniscus islandicus, Macrobiotus crenulatus,
Macrobiotus spectabilis, Macrobiotus willardi, Adorybiotus granulatus, Amphibolus nebulosus,
Rarnazzottius cataphractus and Isobpsibius glaber can be considered typical of tundra
environments, as they have almost never (or very rarely) been found elsewhere.
REFERENCES
Bartog E. 1940. Uber die Variation der art Hypsibius ornatus Richt. (Tardigrada). <oologischJahrbiicher Abteilungjk
Systematik 73: 369-384.
Bernard EC. 1977. A new species of Hexapodibius from North America, with a redescription of Diphascon belgicea
(Tardigrada). Transactcons of the Ammian Mimoscopical Soc&y 96(4): 476482.
Bertolani R, Kristensen RM. 1987. New records of Eohypsibius nadjae and revision of the taxonomic position of
two genera of Eutardigrada (Tardigrada). Biology of Tardigrada, Selected Symposia and Monograph U.<.I:, 1:
359-372.
Bertolani R, Rebecchi L. 1993. A revision of the Macrobiotus hujlandi group (Tardigrada, Macrobiotidae),with
some observations on the taxonomic characters of eutardigrades. xoologica S c r i p 22(2): 127-152.
Dastych H, 1980. Niesporczaki (Tardigrada) Tatrzanskiego Parku Narodowego. Polska W e m i a Nauk, Monopg/ie
Fauny Polski 9: 1-232.
Dastych H. 1983. Two new Tardigrada species from West Spitsbergen and the Tatra Mts. Bulletin de la Sociefk des
Amis des Sciences et des Lettres de Poznam, SeriS 0,ScimCes biologiques 23: 195-200.
Dastych H. 1985. West Spitsbergen Tardigrada. Acta <oologica Crmozimis, Krakow 28(3): 164-214.
Dastych H. 1988. The Tardigrada of Poland. Monogruje F a u v Polski 16: 1-255.
Durante Pasa M V , Maucci W. 1972. Descrizione di Hypsibius (Isohypsibius) basalouoi sp. nov. e altre notizie su
tardigradi del Veronese. M a o h del Muse0 Ciuico di Stork Naturale, Verona 20: 275-281.
Durante Pasa M V , Maucci W. 1975. Descrizione di tre nuove specie di Tardigradi della Scandinavia.Atti deNa
Societu Italiana di S c k z e Naturali e del Muse0 CiliiGo di Storia Naturale di Milano 116(3-4): 244-250.
Durante Pasa M V , Maucci W. 1979. Moss Tardigrada from the Scandinavian Peninsula. Second International
Symposium on Tardkada. Krakaw, Poland, 3 4 28-30 1977, <esz& NaNaukwe Uniwer~tetuJ~lloskiego,Pram <oologiczme,
Krakow 79(25): 47-85.
Grengaard A, Kristensen NM, Petersen MK. 1990. Tardigradfaunaen p i Disco. Feltkiirsus i Arktisk biologi.
Godhavn, 1990 155-179.
Horning DS, Schuster ROYGrigarick AA. 1978. Tardigrada of New Zealand. N m xealandJourna1 o f < o o l o ~ ,
5: 185-280.
It0 M. 1991. Taxonomic study on the Eutardigrada from the northern slope of Mt. Fuji, Central Japan. I. Families
Calohypsibiidae and Eohypsibiidae. Proceedings ofthe Japanese Sockp of systematic <oology 45: 3 M 3 .
Kristensen RM. 1982. New aberrant eutardigrades from homothermic springs on Disko Island, West Greenland.
In: Nelson DR ed, Proceedings ofthe Ihird Intmational symposium on Tardkada. East Tennessee State University
Press, Johnson Cky, Tennessee: 221-236.
Marcus E. 1928. Spinnentiere oder Arachoides. IV Bartierchen (Tardigrada). T m e l t Deutschlands und der
ayenzenden MeeresteileJena 12: 1-230.
Marcus E. 1936. Tardigrada. Das T i i c h 66: 1-340.
Maucci W. 1974. Hypsibius (H.) cataphractus (Tardigrada: Macrobiotidae) und weitere Nachrichten uber
Tardigraden aus Osterreich. Bericht des N a t u n u i s s a f c h a - M e d i z i n ~ cVerins
~
in Innsbluck 61: 83-86.
Maucci W. 1980. Analisi preliminare di alcuni dati statistici sulla ecologia dei Tardigrada muscicoli. Bollettino d d
Muse0 Ciuico di Stork Naturale, Verona 7: 1 4 7 .
Maucci W. 1985. Materiali per una revisione del genere Echiniscw Schultze, 1840. I. I1 complesso blumi
(Heterotardigrada, Echiniscidae). Bollettino del Muse0 Civic0 di Storia Naturale, Verona 12: 104-139.
Maucci W. 1986. Tardigrada. Fauna d’ltalia, vol. 24: Bologna: Calderini, 1-388.
Maucci W. 1988 (1991). Tre nuove specie di Eutardigradi della GroenIandia meridionale.Bollettino delMuseo Ciuio
di Storia Naturale, Vmona 15: 27S289.
Morgan CI. 1980. A systematic survey of Tardigrada from Iceland, Acta Naturalia Zslandica 27: 1-25.
Murray J. 1907. Arctic Tardigrada, collected by Wm. S. Bruce, ‘Transactions ofthe Royal Sociep ~ E d i n b u r g h45(3):
40 1 4 15.
Petersen B. 1951. The tardigrade fauna of Greenland. A faunistic study with some few ecological remarks.
Meddelelser om Grmland, KBbenhavn 150(5): 5-94.
M'.MAUCCI
204
Pilato G , Binda MG. 1977. Precisazioni e rettifiche alla dcscrizione di alcune specie di Tasdiqadi (seconda nota).
Animlia, Catamla 4( I -2): 35-5 1.
pilot0 G , Claxton S, Binda MG. 1989. 'I'ardigrades from Australia. 11. The evaluation of Culohypsibius omatus
(Richtrrs, 1990) caelatus (Marcus, 1928) as a valid species and description of Minibio&r fallax n. sp. (Eutardigrada).
Aninialia 16: 21 -27.
Ramazzotti G, Maucci W. 1983. I1 Philurn 'l'ardigrada. 111 cdizione riveduta ed aggiornata. Mmorie dell'lstituto
Idiano di Idrobiologin, Pallan~a41: 1 1012.
Richters F. 1903. Nordische Tardigaden. <oologisr/w ilnzeiger 27: 168-172.
Richters F. 1904a. Islandische Tardigaden. 300logisch Anzeigu 28(10): 373-377.
Richters R. 1904b. Arktische Tardigraden. Fauna Arclzca 3: 495- 508.
Richters F. 1911a. Faune de Mousses. Tardigades. Camp. arkt. Duc d'&hms, 1907 1-20.
Richters F. 1911b. hloosfauna, in A. Komig, AziJuuna Spit&rgmir, 283-286.
Scourfield DJ. 1897. Contributions to the non-marine Fauna of Spitshergen. Proreedings of thp <oological So&&,
London. 1897 i8+7Y'.
S b & a Y, Elin D. 1989. 1 2 s tardigades du Groenland. RCslutats de la 2eme exptdition du G.E.C.R.P. au
Srorebysund (c6te orientale
198.5). Rrdletin dlensuel C la Sacitti Linnlmne, I:yon, 58(4): 12.1-134.
Srngria Y. 1990. Ixs tardigrades de la cBte orientale du Groenland. Resultats de la troisikme exptdition du
G.E.C.K.P. au Ycorehysiirid ( 1 987). Bulletin Xlmuel dc la SoCiPti Linnimne, Lyon 59(8):325 -332.
Weglarska B. 1965. Die Tardigaden (Tardigrada) Spitzhergcns. dcta <oolngicn CracooienSin, K h w 1l(2):
1:i- 52.
~
~~~