The Tardigrada Fauna of
Łukasz Kaczmarek1 and Łukasz Michalczyk2,*![]()
1 Department of
Animal Taxonomy and Ecology, A. Mickiewicz
University, Umultowska 89, 61-614 Poznań, Poland
E-mail: kacztar@poczta.fm
2 Institute of Environmental Sciences,
E-mail: agnostic@poczta.fm
(Accpeted April 26, 2005)
*Present
address: Centre for
Ecology, Evolution and Conservation,
Łukasz Kaczmarek and
Łukasz Michalczyk (2006) The Tardigrada fauna of
Key words: New species, New records, Taxonomy,
Faunistics.
----------------------------------------------------------------------------------------------------------------
Introduction
The phylum Tardigrada consists of small (50~1200 µm), bilaterally
symmetrical micrometazoans commonly called “water bears”. Tardigrada are divided into 3 classes:
Heterotardigrada (armored), Eutardigrada (unarmored) and Mesotardigrada (with
only 1 dubious species, Thermozodium
esakii Rahm, known from Japanese thermal springs). Currently, more than 800 terrestrial
and/or freshwater species are known (McInnes 1994 and later descriptions).
Papers on Mongolian tardigrades were first published by Iharos (1965
1968 1973) and Maucci (1988). These
papers reported a total of 43 taxa, including: Echiniscus filamentosus
mongoliensis Iharos and E. filamentosus f. aspinosa Iharos which are
now considered synonyms of Echiniscus testudo (Doyére), and Macrobiotus
mongolicus Maucci which was new to science. Another species mentioned in Iharos
(1973) as Hypsibius sp. was an unidentifiable exuvium that could have
been either a Hypsibius or Diphascon. In 1980, Prof. Anna Pacyna (
In this paper 62 species of Mongolian tardigrades are reported. The list is based on earlier papers
published by Iharos (1965 1968 1973) and Maucci (1988), slides made by Prof.
Węglarska, and specimens found in the samples collected from northern
----------------------------------------------------------------------------------------------------------------
MATERIALS AND METHODS
During a scientific expedition to
All measurements of the new species
are given in micrometers (µm).
Structures were measured only if their
orientation was suitable. Body
length was measured from the anterior tip to the end of the body, not including
the hind legs. Buccal tube length
and the level of the stylet support insertion point were measured according to
Pilato (1981). Buccal tube widths
were measured as the external diameters at the level of the stylet support
insertion point. Lengths of the
claw branches were measured from the base of the claw to the top of the branch,
including accessory points. The pt ratio is the ratio of the length of a
given structure to the length of the buccal tube expressed as a percentage
(Pilato 1981). In the description
of the holotype, the pt ratio is
given after the value in micrometers in square brackets and italics ( [12.9] ).
Photomicrographs were made using a phase-contrast microscope (PCM; manufacturer, location) and a Nomarski differential interference contrast (DIC) microscope (give manufacturer, location). All drawings were made using a camera lucida associated with the DIC microscope.
Measurements of I. macrodactylus
(Maucci) and
Numbers of specimens and samples in
which the specimens were found are given in parentheses after the elevation (in
meters above sea level); e.g. (10/4) indicates 10 specimens found in 4 samples,
and (2+5/3) indicates 2 specimens and 5 eggs found in 3 samples).
If the name of a researcher is not
indicated, it means that those specimens were collected and prepared by Ł.
Kaczmarek.
----------------------------------------------------------------------------------------------------------------
RESULTS
List of
species recorded in
a Species found during the present study as new to
b Species found
in
Class Heterotardigrada
Marcus
Order Echiniscoidea Marcus
Family Echiniscidae Thulin
Genus Bryodelphax Thulin
1. Bryodelphax asiaticus Kaczmarek and
Michalczyk 2004 b
Localities in
Genus Cornechiniscus Maucci and Ramazzotti
2. Cornechiniscus
cornutus (Richters)
Cornechiniscus
cornutus
(Richters): Maucci 1988.
Localities in
3. Cornechiniscus holmeni (Petersen)
Cornechiniscus
holmeni
(Petersen): Maucci 1988.
Localities in
4. Cornechiniscus lobatus (Ramazzotti)
Pseudechiniscus lobatus Ram.: Iharos 1973.
Localities in
Genus Echiniscus Schultze
5. Echiniscus baius Marcus
Echiniscus baius Marcus: Iharos 1968.
Localities in
6. Echiniscus blumi Richters a
Localities in
Echiniscus canadensis J. Murr.: Iharos 1973.
8. Echiniscus granulatus (Doyére)
Echiniscus granulatus Doy.: Iharos 1973.
Localities
in
9. Echiniscus
merokensis Richters a
Localities
in
10. Echiniscus reticulatus
Localities
in Mongolia: Arhangaj Ajmag, Terchijn Cagaan Nuur NP: Tarbagataj Mts., mosses from rocks
11. Echiniscus simba Marcus
Echiniscus simba Marcus: Iharos 1968.
Localities in
12. Echiniscus testudo (Doyére)
Echiniscus testudo Doy.: Iharos 1968; E.
filamentosus mongoliensis Iharos, 1973: Iharos 1973; E. filamentosus f. aspinosa Iharos, 1973: Iharos 1973; E. testudo (Doy.):
Maucci 1988.
13. Echiniscus trisetosus Cuénot
Echiniscus trisetosus Cuénot: Iharos 1973.
Localities
in Mongolia: Ömnogöw Ajmak (Iharos 1973); Arhangaj Ajmag, Terchijn
Cagaan Nuur NP: field of lava (with Larix sp. and Pinus sp.
forest) near the dead volcano Chorgijn Togoo (Chorgo), moss from lava,
14. Echiniscus
wendti Richters
Echiniscus wendti Richt.: Iharos 1968.
Genus Pseudechiniscus Thulin
15. Pseudechiniscus jiroveci Bartoš
a
Localities
in Mongolia: Arhangaj Ajmak, east slope of valley near the city
of Cecerleg: mosses from stones,
16. Pseudechiniscus
suillus (Ehrenberg)
Pseudechiniscus suillus Ehrb.:
Iharos 1968.
Localities
in
Genus: Testechiniscus Kristensen
17. Testechiniscus spitsbergensis (Scourfield)
Echiniscus spitsbergensis Scourf.: Maucci 1988.
Localities
in Mongolia: Ömnogöw Ajmak (Maucci 1988); Arhangaj
Ajmak, Terchijn Cagaan Nuur NP: field of lava (with Larix
sp. and Pinus sp. forest) near the dead volcano Chorgijn Togoo (Chorgo),
moss from lava,
Order Parachela Schuster et al.
Family Macrobiotidae Thulin
Genus Macrobiotus Schultze
18. Macrobiotus
areolatus
Macrobiotus areolatus J.
Murr.: Iharos 1968.
Localities
in Mongolia: Töw
Ajmak (Iharos 1968); Arhangaj Ajmak, Terchijn
Cagaan Nuur NP: field of lava (with Larix sp. and Pinus
sp. forest) near the dead volcano Chorgijn Togoo (Chorgo), moss from a wet,
shallow cave,
19. Macrobiotus
harmsworthi
Localities
in Mongolia: Arhangaj Ajmak, east slope of valley near the city
of Cecerleg: mosses from stones and soil,
20. Macrobiotus
hufelandi Schultze
Macrobiotus hufelandii C.A.S. Schultze: Iharos
1965; Macrobiotus hufelandii Schultze: Iharos 1968; Macrobiotus
hufelandii Schultze: Iharos 1973.
Localities
in Mongolia: Chentej Ajmak (Iharos 1968), Töw Ajmak (Iharos 1965 1968);
Ömnogöw Ajmak (Iharos 1973); Arhangaj Ajmak,
east slope of valley near the city of Cecerleg: mosses and lichen from stones,
soil and dead wood
21. Macrobiotus
islandicus Richters
Macrobiotus islandicus Richt.: Iharos 1968.
Localities
in
22. Macrobiotus
mongolicus Maucci b
Macrobiotus mongolicus sp. nov.: Maucci 1988.
Localities
in
23. Macrobiotus
montanus
Localities
in
24. Macrobiotus
richtersi
Macrobiotus richtersi J. Murr.: Iharos 1965; Macrobiotus
richtersi J. Murr.: Iharos 1968; Macrobiotus richtersi J.
Murr.: Iharos 1973.
Localities
in Mongolia: Töw Ajmak (Iharos 1965 1968); Ömnogöw Ajmak (Iharos 1973);
Arhangaj Ajmak, Terchijn Cagaan Nuur NP: field
of lava (with Larix sp. and Pinus sp. forest) near the dead
volcano Chorgijn Togoo (Chorgo), mosses from lava,
25. Macrobiotus spectabilis
Thulin a
Localities
in
26. Macrobiotus willardi Pilato a
Localities
in
Genus Minibiotus Schuster
27. Minibiotus
intermedius (Plate)
Macrobiotus intermedius Plate: Iharos 1968.
Localities
in Mongolia: Töw
Ajmak (Iharos 1965 1968); Arhangaj Ajmak,
east slope of valley near the city of Cecerleg: moss and lichen from stone,
28. Minibiotus
weglarskae Michalczyk, Kaczmarek and Claxton, 2005 b
Localities
in Mongolia: Arhangaj Ajmak, Terchijn Cagaan Nuur NP: field
of lava (with Larix sp. and Pinus sp. forest) near the dead
volcano Chorgijn Togoo (Chorgo), mosses and lichen from lava,
Genus Murrayon Bertolani and Pilato
29. Murrayon hibernicus (
Localities
in
Genus Richtersius Pilato and Binda
30. Richtersius
coronifer (Richters)
Macrobiotus coronifer Richt.: Iharos 1973; Adorybiotus
coronifer (Richters): Maucci 1988.
Localities
in Mongolia:
Ömnogöw Ajmak (Iharos 1973, Maucci 1988);
Arhangaj Ajmak, Terchijn Cagaan Nuur NP: Tarbagataj Mts., mosses from
rocks, 2500 (27+18/1), 2700 ml, (12+3/2), 19 Jul. 2000; Chubsugul Ajmak, Chubsugul NP, Larix sp.
forest near Chubsugul Lake: moss from rock, 1650 (28+20/1),
Family Calohypsibiidae Pilato
Genus Calohypsibius Thulin
31. Calohypsibius
ornatus (Richters)
Hypsibius ornatus Richt.: Iharos 1968.
Localities in
Family Hypsibiidae Pilato
Genus Hypsibius Ehrenberg
32. Hypsibius convergens Urbanowicz
Hypsibius convergens Urb.: Iharos 1965; Hypsibius
convergens Urb.: Iharos 1968.
Localities in Mongolia: Töw Ajmak (Iharos
1965 1968); Arhangaj Ajmak, east slope of valley near the city of
Cecerleg: moss from stone,
33. Hypsibius
dujardini (Doyére)
Hypsibius dujardini Doy.:
Iharos 1968.
Localities
in Mongolia: Töw Ajmak (Iharos 1968); Arhangaj Ajmak, east slope of
valley near the city of Cecerleg: moss from soil,
34. Hypsibius
microps Thulin
Hypsibius microps Thul.: Iharos 1968.
Localities
in
35. Hypsibius novemcinctus Marcus
Hypsibius novemcinctus Marcus: Iharos 1968.
Localities in
36. Hypsibius
pallidus Thulin
Hypsibius pallidus Thul.: Iharos 1968.
Localities
in Mongolia: Töw Ajmak (Iharos 1968); Arhangaj Ajmak, Terchijn Cagaan Nuur NP:
Tarbagataj Mts., moss from rocks,
37. Hypsibius
scabropygus Cuénot a
Localities
in Mongolia: Chubsugul Ajmak, Chubsugul NP, Larix sp. forest near Chubsugl
Lake: lichen from tree
38. Hypsibius sp. (exuvium)
Hypsibius sp.: Iharos 1973.
Localities
in
Genus Isohypsibius Thulin
39. Isohypsibius altai Kaczmarek and
Michalczyk sp. nov.
Holotype: Body length 260.3 (Fig. 1). Body transparent, eyes large, composed
of small black dots. Cuticle smooth on dorsal and ventral sides of body. Bucco-pharyngeal apparatus of Isohypsibius
type (Figs 2, 4). Mouth
anteroventral. Buccal tube 29.5
long and 3.8 [12.9] wide. At
end of buccal tube triangular apophyses present. Stylet supports inserted on buccal tube
at 19.5 [66.1]. Pharyngeal
bulb spherical with 2 macroplacoids and microplacoid. First macroplacoid 5.7 [19.3]
long with central constriction, 2nd one 3.8 [12.9] long. Microplacoid 1.4 [4.7] long. Macroplacoid row 10.5 [35.6]
long. Placoid row 12.4 [42.0]
long. Claws of Isohypsibius
type and very long (Figs. 3, 5).
External claws on leg I: primary branch (pb): 21.9, secondary branch
(sb): 9.5, leg II: pb: 23.8, sb: 9.5, leg III: pb: 26.6, sb: 11.4, leg IV: pb:
31.4, sb: 14.3. Primary branches of
claws without accessory points.
Lunules absent
but bases of claws expanded (Figs 3, 5).
Cuticular bars on 1st 3 pairs of legs present near base of claw (Fig.
1). Eggs white, smooth, and
deposited in exuvium.

Figs. 1-3. Isohypsibius altai sp. nov. (1) Habitus; (2) buccal apparatus; (3)
claws of the 4th pair of legs (DIC).

Figs. 4, 5. Isohypsibius altai sp. nov. (4 ) Buccal apparatus; (5) claws of the
4th pair of legs.
Etymology
The
name ‘altai’ refers to the name of the Altai Mts. in
Material examined:
Holotype and 5 paratypes: Southern Mongolia,
Additional material:
One specimen
in the simplex stage and 4 exuvia with eggs.
Type depositories: Holotype is deposited at the Zoological Museum of
Jagiellonian University, Ingardena 6, 30-060
Differential diagnosis:
Isohypsibius altai sp. nov. is similar to I. macrodactylus and
40. Isohypsibius archangajensis
Kaczmarek and Michalczyk b
Localities in
41. Isohypsibius
mihelcici (Iharos)
Hypsibius mihelcici Iharos: Iharos 1965; Hypsibius
mihelcici Iharos: Iharos 1968.
Localities
in Mongolia: Töw Ajmak (Iharos 1965 1968); Arhangaj Ajmak, east slope of valley near
the city of Cecerleg: mosses from soil,
42. Isohypsibius
nodosus (
Localities
in
43. Isohypsibius
sattleri (Richters)
Hypsibius bakonyiensis Iharos: Iharos 1965; Hypsibius
bakonyiensis Iharos: Iharos 1968.
Localities
in Mongolia: Töw Ajmak (Iharos 1965 1968); Arhangaj Ajmak, east slope of valley near
the city of Cecerleg: mosses from stones,
44. Isohypsibius
schaudinni (Richters)
Hypsibius schaudinni Richt.: Iharos 1968; Hypsibius
schaudinni Richt.: Iharos 1973.
Localities
in
45. Isohypsibius
tuberculatus (Plate)
Hypsibius tuberculatus Plate: Iharos, 1965; Hypsibius
tuberculatus Plate: Iharos, 1968.
Localities
in
Genus Thulinius Bertolani
46. Thulinius
augusti (
Hypsibius augusti J.
Murr.: Iharos 1968.
Localities
in
Remarks: The taxonomic status of Hypsibius augusti is still unclear
(Bertolani et al. 1999, Nelson et al. 1999) and without examination of the
material examined by Iharos, it is impossible to decide to which genus it
should be attributed. At this stage
we have decided to attribute this species to the genus Thulinius, but future research is required on this matter.
Genus Astatumen Pilato
47. Astatumen ramazzotti (Iharos)
Itaquascon ramazzotti Iharos: Iharos 1968.
Localities
in
This species is probably a synonym of A. trinacriae (Arcidiacono).
48. Astatumen trinacriae (Arcidiacono) a
Localities
in Mongolia: Arhangaj Ajmak, Terchijn Cagaan Nuur NP: field of lava (with Larix
sp. and Pinus sp. forest) near the dead volcano Chorgijn Togoo (Chorgo),
moss from lava,
Genus Itaquascon Barros
49. Itaquascon
mongolicus Kaczmarek, Michalczyk and Węglarska, 2002 b
Localities
in Mongolia: Arhangaj Ajmak, Terchijn Cagaan Nuur NP: Tarbagataj Mts., moss
from rock,
Genus Diphascon Plate
Subgenus
Diphascon Pilato
50. Diphascon (D.) brevipes (Marcus)
Diphascon brevipes (Marcus): Maucci
1988.
Localities
in
51. Diphascon
(D.) bullatum
Hypsibius bullatus J. Murr.: Iharos 1968.
Localities
in
52. Diphascon (D.) pingue (Marcus)
Hypsibius pinguis Marcu: Iharos 1965; Hypsibius
pinguis Marcus.: Iharos 1968.
Localities
in Mongolia: Töw Ajmak (Iharos 1965 1968); Arhangaj Ajmag, Terchijn
Cagaan Nuur NP: field of lava (with Larix sp. and Pinus sp.
forest) near the dead volcano Chorgijn Togoo (Chorgo), moss from a wet, shallow
cave,
53. Diphascon
(D.) recamieri Richters
Hypsibius recamieri Richt.: Iharos 1965; Hypsibius
recamieri Richt.: Iharos 1968.
Localities
in
54. Diphascon (D.) sexbullatum Ito
(Figs. 6-12) a
Localities
in Mongolia: Arhangaj Ajmak, east slope of valley near the city of Cecerleg: mosses
from stones,
This is a 2nd record of this species. We also provide here for the very 1st
time photomicrographs of the holotype of this species.

Figs 6, 7. Diphascon sexbullatum; habitus. (6 ) Specimen from

Figs. 8-12. Diphascon sexbullatum. (8) Dorsal cuticle with gibbosities; (9,
10) buccal apparatus; (11, 12) claws of the 4th pair of legs (8, 10, and 12:
holotype, PCM; 9 and 11: a specimen from
Subgenus Adropion Pilato
55. Diphascon (A.) belgicae Richters
Hypsibius belgicae Richt.: Iharos 1968.
Localities
in
56. Diphascon (A.) prosirostre Thulin
Hypsibius prosirostris Thul.: Iharos 1968.
Localities
in
57. Diphascon (A.) scoticum
Hypsibius scoticus J. Murr: Iharos 1968.
Localities
in
Genus Hebesuncus Pilato
58. Hebesuncus
conjugens (Thulin) a
Localities
in
Genus Platicrista Pilato
59. Platicrista
horribilis Kaczmarek and Michalczyk b
Localities in
Genus Ramazzottius Binda and Pilato
60. Ramazzottius
oberhaeuseri (Doyére)
Hypsibius oberhaeuseri Doy.: Iharos 1968; Hypsibius
oberhaeuseri Doy.: Iharos 1973.
Localities in
Mongolia:
Töw Ajmak (Iharos 1968); Ömnogöw Ajmak (Iharos 1973); Arhangaj
Ajmag, Terchijn Cagaan Nuur NP: field of lava (with Larix sp. and
Pinus sp. forest) near the dead volcano Chorgijn Togoo (Chorgo), moss
from lava,
Order Apochela Schuster et al.
Genus Milnesium Doyére
61. Milnesium
cf. eurystomum Maucci a
Localities
in
62. Milnesium
tardigradum Doyére
Milnesium tardigradum Doy.: Iharos 1965; Milnesium
tardigradum Doy.: Iharos 1968; Milnesium tardigradum Doy.: Iharos
1973; Milnesium tardigradum Doy.: Maucci 1988.
Localities in
Mongolia:
Töw Ajmak (Iharos 1965 1968); Ömnogöw Ajmak (Iharos 1973, Maucci
1988); Arhangaj Ajmag, Terchijn Cagaan Nuur NP: field of lava (with Larix
sp. and Pinus sp. forest) near the dead volcano Chorgijn Togoo (Chorgo),
moss from lava,
A
key to species reported from
1. Dorsal side of body covered with
cuticular plates, lateral cirrus A present .....……............. 2
- Dorsal side of body without plates,
lateral cirrus A absent ..................................……......... 18
2. Pseudosegmental plate present
..............……………………………………………..…...... 3
- Pseudosegmental plate absent
........................................................................……….............
7
3. Cirri A in the shape of
filaments ……………………………………………...…………… 4
-. Cirri A in the shape of
short, thick cones …………………………………...………….. 5
4. Between scapular and median plate I 4 additional plates present
……………….....……..
...................................................................................................
Pseudechiniscus jiroveci Bartoš
-. Additional plates absent
................................................ Pseudechiniscus suillus (Ehrenberg)
5. Besides cirri A, other lateral or dorsal
filaments, spines, or teeth absent ………………… 6
-. Lateral filaments C and D, dorsal
spine Dd, and
teeth on the margin of pseudosegmental plate present
…………………………………………...… Cornechiniscus holmeni (Petersen)
6. Posterior margin of pseudosegmental plate
smooth, large triangular teeth absent from pair IV of legs……………………………………………….… Cornechiniscus
cornutus (Richters)
-. Posterior margin of pseudosegmental plate with
2 lobes or teeth, large triangular teeth on pair IV of legs present
……………………………….. Cornechiniscus lobatus (Ramazzotti)
7. Median plates I and II not divided, incisions
on terminal plate present ……...……….. 8
-. Median plates I and II divided, incisions on terminal plate absent
……………………….
……………………………..……. Bryodelphax asiaticus Kaczmarek and Michalczyk (in press)
8. Two rows of ventral plates absent
……………………………………………………...….. 9
-. Two rows of ventral plates present
………..…..... Testechiniscus spitsbergensis (Scourfield)
9. Lacking
any other lateral or dorsal appendages than cirrus A …………………....…..… 10
-. Lateral
or/and dorsal appendages other than cirrus A present ……….…………...….….. 11
10. Median plate III present, terminal
plate facetted, smooth transverse band absent from anterior part of paired plates I and
II ……….…………..…… Echiniscus reticulatus
-. Median plate III absent, terminal plate not
facetted, smooth transverse band present in anterior part of paired plates I
and II ……….................………… Echiniscus
wendti Richters
11. Dorsal appendages (spines and/or
filaments) absent .................... Echiniscus baius Marcus
-. Dorsal appendages (spines and/or
filaments) present ………………………………..…… 12
12. Lack of
any other lateral appendages than cirrus A ……..... Echiniscus canadensis
-. Lateral
appendages other than cirrus
A present
…….................………………………….. 13
13. Appendages
E present ………………………………………...…………………………
14
-. Appendages
E absent or with very small
difficult-to-see spine ………..…………….… 16
14. All
lateral appendages as filaments ……………………………...…………………….. 15
-. Lateral
appendages B, D, and E as spines
…………………….…. Echiniscus simba Marcus
15. Appendage
B absent, appendage D present
…………...…. Echiniscus merokensis Richters
-. Appendage
B present, appendage D absent
……........................ Echiniscus
testudo (Doyére)
16. Appendages B present ……………………………………..……………………………. 17
-. Appendages B absent …………………………………..……... Echiniscus trisetosus Cuénot
17. Dorsal appendages C
as filaments, spine E absent,
external claws on IV pair of legs with 1~3 spurs ……………………………………………………..…..... Echiniscus blumi Richters
-. Dorsal appendages C
as thick spines, very small spine E
present, internal claws on pair IV of legs with spurs ………………………..……………..….… Echiniscus
granulatus (Doyére)
18. Peribuccal and cephalic
papillae absent …………………………….……….………….. 19
-. Peribuccal and cephalic
papillae present ………………………………………………..... 61
19. Claws symmetrical;
external and internal claws identically developed (of similar size and shape)
………………………………………………………………………..…………… 20
-. External and internal
claws always more or less dissimilar in shape and/or size
…………………...…………………………………………………………………………... 32
20. Thickness of buccal tube
wall decreasing from posterior to anterior part ……….………....
……………………………………………………………..…. Richtersius coronifer (Richters)
-. Thickness of buccal tube wall not
decreasing from posterior to anterior part …………..... 21
21. Peribuccal lamellae present,
peribuccal papulae absent, stylet supports inserted at more than 73% of buccal
tube length, buccal tube wide (> 12% of buccal tube length) …………….... 23
- Peribuccal lamellae absent,
peribuccal papulae present, stylet supports inserted at 54% of buccal tube
length, buccal tube narrow (≤ 6.8% of buccal tube length)
......................... 22
22. Cuticle smooth without
pores ............................................... Minibiotus intermedius (Plate)
-. Cuticle with distinct
pores ……………………...……………………...…..............................
............................................. Minibiotus weglarskae Michalczyk, Kaczmarek and Claxton, 2005
23. Claws of pullari type,
cuticle with distinct areas of evident granulation on lateral side of body
……………………...……………………………..… Murrayon hibernicus (
Claws of hufelandi type, cuticle without granulations on lateral
side of body ……….….. 24
24. Cuticle with pores
………………………………………….…………………………… 25
-. Cuticle without pores
……………………………………………………….…………….. 29
25. Pharynx with 2
macroplacoids …………………………………….……………………. 26
-. Pharynx with 3
macroplacoids …………………..………………….…………….………. 28
26. Microplacoid present
……………………………………………………………………. 27
-. Microplacoid absent
…………………………..…………... Macrobiotus
islandicus Richters
27. Eggs of hufelandi type with inverted goblet-shaped processes……………….…………….
…………………………………………………………..…… Macrobiotus hufelandi Schultze
-. Eggs of tenuis type with processes in the shape
of truncated cones ……………....………….
……………………………………………………………..... Macrobiotus mongolicus
Maucci
28. Pores present on entire
cuticle, egg surface between processes with areolation, processes in the shape
of truncated cones …..…………….……………… Macrobiotus
richtersi
-. Pores present only on
legs, egg surface between processes without areolation, processes in the shape
of flexible cones ……………………………………….. Macrobiotus
willardi Pilato
29. Pharynx with 2
macroplacoids ………...…...…………..… Macrobiotus
spectabilis Thulin
-. Pharynx with 3
macroplacoids …………………..………………….…………….………. 30
30. Microplacoid present,
egg surface between processes without areolation, macroplacoids rounded
……...................................................................................................................…….
31
-. Microplacoid absent, egg
surface between processes with areolation, macroplacoids elongated
……………………………………………..……….. Macrobiotus
areolatus
31. Egg processes in shape
of cones with reticular design on surface, surface between processes without
punctuation ………………...………..… Macrobiotus harmsworthi
-. Egg processes in the
shape of small hemispheres, surface between processes with punctuations which
form a reticular design …..………….……Macrobiotus montanus
32. Claws of Calohypsibius type, less then 5 µm long,
dorsal side of the body covered by spines arranged in transverse bands
…………................….. Calohypsibius
ornatus (Richters)
-. Claws of different type,
more then 5 µm long, dorsal
side of body with tubercles/gibbosities
..................................................................................................................................................
33
33. Claws of Isohypsibius type, external and internal claws similar in shape and usually differing in size …………………………………………….....………………..…………. 34
-. Claws of different type,
external and internal claws differing in size and shape ……..….. 41
34. Twelve peribuccal lamellae present, 3rd
band of teeth present, freshwater species …..
………………………………………….…………..………..……. Thulinius augusti (
-. Twelve peribuccal lamellae absent, 3rd band
of teeth absent, terrestrial species ..….. 35
35. Dorsal cuticle without gibbosities ……………………………………….……………… 36
-. Dorsal cuticle with gibbosities arranged in transverse rows ………………...……..……. 38
36. Cuticle smooth, without
undulations ………………………….…………………….. 37
-. Cuticle on dorsal side
of body with rounded depressions forming reticular design and with 14~16
undulations ………………… Isohypsibius
archangajensis Kaczmarek and Michalczyk
37. Pharynx with 3
macroplacoids, claws on pair IV of legs ~20
µm long ……………….
………………………………..………………..………….. Isohypsibius schaudinni (Richters)
-. Pharynx with 2
macroplacoids, claws on pair IV of legs 30.4~42.8 µm long ………………..
………………………………………………………………...…… Isohypsibius altai sp. nov.
38. All rows
containing even number of gibbosities ……………………….………………..39
-. Some rows
containing odd number of gibbosities …………………………………...… 40
39. Gibbosities
arranged in 7 transverse rows: rows 1~5 with 6 gibbosities, rows 6 and 7 with 4
……………………………………………………..……. Isohypsibius nodosus (
-.
Gibbosities arranged in 9 transverse rows with 2, 4, or 6 gibbosities; tubercles/gibbosities with 2 or 3
short spines at tips ……... Isohypsibius sattleri (Richters)
40. Gibbosities
arranged in 10 transverse rows: 1st and 10th rows with 5 gibbosities; 2nd, 4th,
6th, and 8th rows with 4; 3rd, 5th, and 7th rows with 6; and 9th row with 2
gibbosities …
………………………………………………………….…... Isohypsibius tuberculatus (Plate)
-. Gibbosities
arranged in 8 transverse rows: 7th row with 2 gibbosities; 2nd, 4th, and 6th
rows with 4; 1st, 3rd, 5th, 8th rows with 3 ………………….... Isohypsibius mihelcici (Iharos)
41. Claws of Ramazzottius type, main branches of external claws completely
separated from rest of claw ………………………………..……. Ramazzottius oberhaeuseri
(Doyére)
-. Claws of Hypsibius type, main branches of external claws
connected with rest of claw ………………………………………………………………………………………..... 42
42. Buccal
tube rigid without spiral thickenings ……………………………………………. 43
-. Buccal
tube divided into rigid anterior part (buccal tube) and flexible posterior part
(pharyngeal tube) with spiral thickenings
...............................................................................
49
43. Cuticle
on dorsal side of body smooth ……………..……..…………………….. 46
-. Cuticle on
dorsal side of body sculptured ……………………...…………………. 45
45. Cuticle
on dorsal side of caudal end body covered by irregular plates, eggs with tiny
granulations ………………………….............……… Hypsibius
scabropygus Cuénot
-. Cuticle
on dorsal side with 9 transverse bands of pigmentation, eggs smooth
…………………………………………………………..…… Hypsibius novemcinctus Marcus
46. Septula present
…………………………………..…………. Hypsibius dujardini (Doyére)
-. Septula absent
…………………………………………………………………………….. 47
47. Macroplacoids rod-shaped, clearly longer then
wide, 1st macroplacoid with a constriction in middle, small transversal bar
between claws on pair IV of legs present, eyes usually absent ………….…………………………………..……… Hypsibius
convergens Urbanowicz
- Macroplacoids rounded or granular,
small transverse bar on pair IV of legs absent, large eyes present
.............................................................................................................................
48
48. Basal portions of claws the same length or
longer than secondary branches
..............
.............................................................................................................Hypsibius
pallidus Thulin
-. Basal portions of claws clearly shorter than
secondary branches) ….................................
…………………………………………………………………..….. Hypsibius microps Thulin
49. Pharynx with 1 very slender macroplacoid
................................................................... 50
- Pharynx with more than 1 macroplacoid
.......................................................................... 52
50. Stylet supports present,
dentate lunules present
.....................................................................
………………………..…….... Itaquascon mongolicus Kaczmarek,
Michalczyk and Węglarska
-. Stylet supports absent, dentate lunules absent
…………………………………...……….. 51
51. Pharynx elongated oval,
claws long and slender ……….… Astatumen ramazzotti (Iharos)
-. Pharynx oval, claws
short and massive ………….…… Astatumen
trinacriae (Arcidiacono)
52. Apophyses for the insertion of
stylet muscles in the shape of ‘blunt hooks’,
pharyngeal tube shorter than buccal tube .………………………….……. Hebesuncus conjugens (Thulin)
-. Apophyses for the insertion of stylet muscles
shaped differently, pharyngeal tube longer than buccal tube ...………………………………………………………………….……….. 53
53. Apophyses for insertion of stylet
muscles in shape of ‘very wide and flat
ridges’, buccopharyngeal tube short and wide ………. Platicrista
horribilis Kaczmarek and Michalczyk
-. Apophyses for insertion of stylet muscles in the shape of ‘semilunar hooks’, buccopharyngeal
tube very long and narrow ………….……………………….…………… 54
54. Drop-like structure between buccal and
pharyngeal tube present .................................... 55
-. Drop-like structure between buccal
and pharyngeal tube absent ........................................ 59
55. Cuticle on dorsal side of body
smooth …………………………..…………….... 56
-. Cuticle on dorsal side of body with
gibbosities covered with reticular design ...…… 58
56. Pharynx with 2 macroplacoids and
septula or microplacoid …………………………… 57
-. Pharynx with 3 macroplacoids,
septula, and microplacoid ……………….………………….
…………………………………………………..…..
Diphascon (Diphascon) pingue (Marcus)
57. Septula present …………………………..…… Diphascon (Diphascon) brevipes
(Marcus)
-. Microplacoid present ……………………..…… Diphascon (Diphascon)
recamieri Richters
58. Gibbosities arranged in 9 transverse
rows with 2 gibbosities in each row ……..………..
………………………………………………..……. Diphascon (Diphascon) bullatum
-. Only 1
longitudinal row with 6 gibbosities …………………………………
…………………………………………..…………… Diphascon (Diphascon)
sexbullatum Ito
59. Pharynx with 2
macroplacoids ……………..…... Diphascon (Adropion) belgicae Richters
-. Pharynx with 3
macroplacoids …………………………………………………...……….. 60
60. Microplacoid and septula
absent ………..…….. Diphascon (Adropion) prosirostre Thulin
-. Microplacoid and septula
present ……………….... Diphascon
(Adropion) scoticum
61. Buccal tube decreasing
in width from anterior to posterior part ……………………
…………………………………………………………..… Milnesium cf. eurystomum Maucci
-. Buccal tube not
decreasing in width from anterior to posterior part …………………
………………………………………………………………....Milnesium
tardigradum Doyére
Acknowledgements:
The
authors want to thank Prof. Barbara Węglarska (
The
paper is based on a Master’s thesis by the 1st author: „Niesporczaki (Tardigrada)
Mongolii” (‘Water bears (Tardigrada) of Mogolia’). The thesis was successfully defended in
June 2003 and is available at A. Mickiewicz University (Department
of Animal Taxonomy and Ecology, A. Mickiewicz
University, Umultowska 89, 61-614
----------------------------------------------------------------------------------------------------------------
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