Zootaxa 2781: 29–39 (2011) ISSN 1175-5326 (print edition)

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Copyright © 2011 · Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)

New records of Tardigrada from Bulgaria with the description of

Macrobiotus binieki sp. nov. (Eutardigrada: Macrobiotidae) and
a key to the species of the harmsworthi group

ŁUKASZ KACZMAREK1, BARTŁOMIEJ GOŁDYN2, ZOFIA M. PROKOP3 & ŁUKASZ MICHALCZYK4

1
Department of Animal Taxonomy and Ecology, A. Mickiewicz University, Umultowska 89, 61-614 Poznań, Poland.
E-mail: kaczmar@amu.edu.pl
2
Department of General Zoology, A. Mickiewicz University, Umultowska 89, 61-614 Poznań, Poland. E-mail: glodny@amu.edu.pl
3
Molecular and Behavioural Ecology Group, Institute of Environmental Sciences, Jagiellonian University, Gronostajowa 7, 30-397
Kraków, Poland. E-mail: zofia.prokop@uj.edu.pl
4
Centre for Ecology, Evolution and Conservation, School of Biological Sciences, University of East Anglia, Norwich NR4 7TJ, UK.
E-mail: LM@tardigrada.net

Abstract

Fifteen moss samples collected in the Sophia Province (Bulgaria) were examined. In these samples six eutardigrade spe-
cies were found: Hypsibius convergens, Isohypsibius prosostomus, Macrobiotus binieki sp. nov., M. hufelandi, M. pallarii
and Ramazzottius oberhaeuseri. The new species belongs to the Macrobiotus harmsworthi group and differs from the most
similar M. australis, M. coronatus, M. patiens, M. pseudocoronatus, M. radiatus, M. rigidus and M. simulans mainly by:
egg processes covered by small bubbles (not smooth or reticulated), egg shell between processes covered by wrinkles not
dots or stripes forming a large radiate crown, a higher number of processes on egg circumference and some morphometric
characters of adults. In this paper a key to all species of the harmsworthi group is also given.

Key words: Tardigrades, Europe, taxonomy, fauna

Introduction

So far only about thirty four tardigrade species (eleven Heterotardigrada and twenty three Eutardigrada) have been
recorded from Bulgaria (McInnes 1994), the majority reported by Iharos (1961; 1973b; 1982) with most being con-
sidered cosmopolitan. This limited literature for the Bulgarian limno-terrestrial tardigrades indicates the need for
further research. We were able to collect samples from the Sofia City Province in western Bulgaria. Situated in the
central Balkans, Sofia city is the capital of Bulgaria and is surrounded on all sides by mountains. The city nestles at
the northern base of Vitosha, a mountain massif well known for hiking and skiing, and home to the oldest national
park in the Balkans. This region, Vitosha Mountains, was sampled by Iharos (1961; 1982) where he reported 19
taxa but none belonging to the Macrobiotus harmsworthi group.
The new species described in this paper, Macrobiotus binieki sp. nov., belongs to the Macrobiotus harmsworthi
group, which is characterised by having three macroplacoids in the shape of short, rounded rods and a microplacoid
situated very close to them, and conical or hemispherical egg processes. Up to now thirty nine species and one sub-
species were described in this group (Ramazzotti & Maucci 1983; Pilato & Binda 2001; Michalczyk & Kaczmarek
2003; Pilato et al. 2004, 2006, Tumanov 2005; Pilato & Lisi 2006a,b, 2009a; Kaczmarek et al. 2007; Rossi et al.
2009). In this paper, we present the results of our study with the description of M. binieki sp. nov., and provide a
key to all known species of the group.

Accepted by S. McInnes: 8 Jan. 2011; published: 2 Mar. 2011 29

Material and methods

Fifteen moss samples were collected by the second author from the Sofia Province on the 27th of August 2007. In
nine samples 133 specimens and 31 eggs were found. These positive samples were collected from the following
locations:

A. Vitosha Mts., ca. half way between the Cherni Vrah and the Sedloto shelter, 2150 m asl, 42°33’47” N,
23°16’08” E, moss from stone.
B. Vitosha Mts., near the entrance to the Sedloto shelter, 2080 m asl, 42°33’56.5” N, 23°15’29” E, moss from
stone.
C. Vitosha Mts., near the Koniarnika ski lift, 1980 m asl, 42°34’42” N, 23°14’52” E, moss from stone.
D. Vitosha Mts., near Koniarnika, forest upper limit, 1880 m asl, 42°34’57” N, 23°14’46.5” E, moss from stone.
E. Vitosha Mts., ca. half way between Koniarnika and the Kumata shelter, 1770 m asl, 42°35’27” N, 23°15’06”
E, moss from soil.
F. Vitosha Mts., near the Kumata shelter, 1720 m asl, 42°35’41” N, 23°15’01” E, moss from wood.
G. Vitosha Mts., near route from the Kumata shelter to Zlatni Mostove, 1500 m asl, 42°36’27” N, 23°14’31” E,
moss from stone.
H. Vitosha Mts., top of the Cherni Vrah, 2290 m asl, 42°33’50” N, 23°16’33” E, moss from stone.
I. Sofia City, Zhitnitsa Street, 600 m asl, 42°40’56” N, 23°16’24” E, moss from pavement.

All specimens were mounted on microscopic slides in Hoyer’s medium. Photomicrographs were taken using
Phase Contrast Microscopy (PCM). All measurements are given in micrometers [µm]. Body length was measured
from the mouth to the end of the body excluding the hind legs. Buccal tube length and level of the stylet support
insertion point were measured from anterior margin of stylet sheaths. Buccal tube widths were measured as the
external and internal diameters at the level of the stylet support insertion point. 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). Terminology describ-
ing the oral cavity armature is given according to Michalczyk & Kaczmarek (2003) and classification of oral cavity
armature types according to Pilato (1972). Species were determined mainly on the basis of the key to the World
Tardigrada (Ramazzotti & Maucci 1983) and original descriptions. All measurements and ratios used in the key
and differential diagnosis were taken or calculated using original species descriptions.

Results

Taxonomic accounts of species found in the study

Macrobiotus hufelandi hufelandi Schultze, 1833

Localities: A: 13 specimens and 7 eggs, B: 3 specimens, C: 2 specimens, D: 12 specimens, E: 7 specimens, F: 1
specimen, G: 3 specimens.
Previously reported from this region by Iharos (1961).

Macrobiotus pallarii Maucci, 1954

Locality: H: 43 specimens and 20 eggs.
Previously not reported from Bulgaria.

Hypsibius convergens (Urbanowicz, 1925)

Locality: F: 1 specimen.
Previously reported from this region by Iharos (1961).

Ramazzottius oberhaeuseri (Doyère, 1840)

Locality: E:10 specimens.
Reported from Bulgaria (Iharos, 1961), but not the environs of Vitosha.

30 · Zootaxa 2781 © 2011 Magnolia Press KACZMAREK ET AL.

Isohypsibius prosostomus Thulin, 1928
Locality: I: 25 specimens.
Previously not reported from Bulgaria.

Macrobiotus binieki sp. nov.

FIGURES 1–8, TABLES 1–2

Material examined: Holotype and 16 paratypes (12 adults + 4 eggs (including two with embryos) were extracted
from a moss sample collected from location A (see Material and Methods).
Description. Adult (measurements in Table 1): Body transparent/white (Fig. 1). Eyes present. Cuticle
smooth, without pores. Fine, regular granulation present on all legs, developed better on hind legs.

TABLE 1. Measurements [in µm] of selected morphological structures of specimens of Macrobiotus binieki sp. nov. mounted
in Hoyer’s medium (RANGE refers to the smallest and the largest structure found among all measured specimens (including
the holotype); N—number of specimens/structures measured, SD—standard deviation).
CHARACTER N RANGE MEAN SD Holotype
µm pt µm pt µm pt µm pt
Body 6 357–647 895–1091 446 965 108 70 379 924
Buccal tube 6 37.2–59.3 – 46.0 – 8.5 – 41.0 –
Stylet support insertion point 6 28.6–46.1 74.9–77.8 35.1 76.4 6.6 1.3 31.9 77.8
Buccal tube external width 6 4.6–7.8 12.0–13.2 5.7 12.4 1.2 0.5 4.9 12.0
Buccal tube internal width 6 3.4–5.8 8.3–9.8 4.1 8.8 0.9 0.6 3.5 8.5
Ventral lamina 6 23.7–39.7 63.2–66.9 29.6 64.1 6.1 1.4 25.9 63.2
Macroplacoid 1 6 5.3–8.5 11.8–16.1 6.2 13.6 1.2 1.5 5.3 12.9
Macroplacoid 2 6 3.0–5.3 7.6–8.9 3.9 8.4 0.9 0.6 3.3 8.0
Macroplacoid 3 6 4.2–7.4 10.2–12.5 5.1 11.0 1.3 0.9 4.4 10.7
Microplacoid 6 2.4–4.4 5.5–7.4 2.9 6.2 0.8 0.7 2.4 5.9
Macroplacoid row 6 14.3–24.2 34.4–40.8 17.1 37.1 3.8 2.4 14.9 36.3
Placoid row 6 17.3–29.4 42.3–49.6 20.9 45.2 4.7 2.8 17.9 43.7
Claw 1 - primary branch 5 8.3–12.5 20.2–22.6 9.5 21.4 1.8 1.1 8.3 20.2
Claw 1 - secondary branch 3 5.9–10.6 14.4–17.9 7.9 16.2 2.4 1.7 5.9 14.4
Claw 2 - primary branch 4 8.3–12.8 20.2–24.0 10.1 22.2 2.1 1.6 8.3 20.2
Claw 2 - secondary branch 3 5.9–9.3 15.7–18.6 7.8 16.7 1.7 1.6 ? ?
Claw 3 - primary branch 3 8.5–13.2 22.3–23.7 10.5 22.9 2.4 0.7 ? ?
Claw 3 - secondary branch 2 6.5–11.4 17.5–19.2 9.0 18.3 3.5 1.2 ? ?
Claw 4 - primary branch 5 10.6–15.3 24.6–31.7 12.8 27.5 1.7 3.0 10.6 25.9
Claw 4 - secondary branch 5 7.7–11.4 17.5–25.0 9.5 20.6 1.3 3.1 7.7 18.8

Bucco-pharyngeal apparatus of the Macrobiotus-type (Fig. 2). Mouth antero-ventral, surrounded by a ring of
10 peribuccal lamellae. Oral cavity armature of the harmsworthi-type, with three well developed bands of teeth.
Teeth of the first band are smaller than those of the other two bands and are visible in PCM as granules. They are
present in the anterior portion of the oral cavity just behind the peribuccal lamellae. This band of teeth is continu-
ous and looks the same on all oral cavity walls. Teeth in the second band are intermediate in size between those of
the first band and those of the third band of teeth. They are in the shape of small granules and ridges parallel to the
main axis of the buccal tube. They are positioned in the posterior portion of the oral cavity just behind the ring fold
and just before the third band of teeth. This band is continuous and teeth are arranged in one row. Teeth of the third
band are larger than those in the other two bands. They are in the shape of transverse ridges/baffles or granules
(PCM). They are positioned in the rear of the oral cavity just behind the second band of teeth and just before the

MACROBIOTUS BINIEKI SP. NOV. Zootaxa 2781 © 2011 Magnolia Press · 31

FIGURES 1–3. Macrobiotus binieki sp. nov. adult: 1. habitus (ventral view, holotype), 2. buccal apparatus (ventral view, holo-
type), 3. claws III.

32 · Zootaxa 2781 © 2011 Magnolia Press KACZMAREK ET AL.

FIGURES 4–8. Macrobiotus binieki sp. nov. egg: 4. mid-section, 5–6. egg surface (at two different focusing levels), 7–8. pro-
cesses. Scale on 6 same as on 5 and scale on 8 same as on 7.

buccal tube opening. Usually this band is not continuous and is divided into two series: ventral and dorsal. Both
series consist of one median and two lateral teeth. The medio-ventral tooth may be broken into two (or more)
smaller teeth, thus there may be seven (or more) teeth in this band.
Buccal tube is strengthened with the ventral lamina and one bend in anterior part of tube (visible in lateral
view). Pharyngeal bulb slightly oval, with apophyses, three macroplacoids and a microplacoid. Pharyngeal apo-
physes distinct, rounded and forked posteriorly. First macroplacoid rod-shaped and thinner anteriorly, second round
or oval, and third rod-shaped with a subterminal constriction. Second macroplacoid placed slightly closer to the
first than to the third macroplacoid. Macroplacoid length sequence (from smallest to largest): 2-3-1. Microplacoid
small, thin and placed close to the third macroplacoid.
Claws of the hufelandi-type, stout (Fig. 3). Primary branches with distinct accessory points. Lunules on all legs
smooth. Thin bars under claws I–III present. Other cuticular thickenings on legs absent.
Egg (measurements in Table 2): White/transparent, laid freely (Figs 4–8). Spherical, without areolation. Pro-
cesses in the shape of long, smooth flexible spines, with very wide bases (or, in other words, very short cones with
extremely long and flexible spines) (Figs 7–8). Processes composed of two walls (internal and external). The two
walls are supported by a system of partitions. Under PMC these partitions are visible as a reticular design for most
species of the harmsworthi group, however, in M. binieki sp. nov. the partitions are thick and as a consequence
seen as small bubbles rather than a mesh (Fig. 5). At the process base there is a crown of rectangular swellings

MACROBIOTUS BINIEKI SP. NOV. Zootaxa 2781 © 2011 Magnolia Press · 33

(which could also be interpret as very small finger-like structures) (Figs 5–6). Processes are distributed on the sur-
face of the egg very close one to another (occasionally in contact). Surface between processes is covered by an
irregular pattern (wrinkles) visible in PCM as darker and lighter areas (Figs 5–6).

TABLE 2. Measurements [in µm] of selected morphological structures of eggs of Macrobiotus binieki sp. nov. mounted in
Hoyer’s medium (RANGE refers to the smallest and the largest structure found among all measured eggs/structures; N—num-
ber of eggs/structures measured, SD—standard deviation).

CHARACTER N MIN MAX MEAN SD

Diameter of egg without processes 3 85.1 94.5 89.6 4.7
Diameter of egg with processes 3 108.7 114.7 112.2 3.1
Processes height 12 9.8 14.5 12.3 1.4
Processes base width 12 6.5 9.0 7.7 0.8
Process base/height ratio 12 0.50 0.79 0.63 0.09
Distance between processes 12 1.9 3.1 2.6 0.4
Number of processes on egg the circumference 3 27 32 29.3 2.5

Type depositories. Holotype and 16 paratypes (12 adults and 4 eggs) are preserved at the Department of Ani-
mal Taxonomy and Ecology, A. Mickiewicz University, Poznań.
Etymology. The new species is named after Janusz Biniek, a mentor of contract bridge players in the city of
Poznań, on the occasion of his 84th birthday.
Differential diagnosis. M. binieki sp. nov. is most similar, in regard to the non-areolated egg shell and a crown
of thickenings around egg processes, to M. australis Pilato & D'Urso, 1976, M. patiens Pilato, et al., 2000, M.
pseudocoronatus Pilato et al., 2006, M. rigidus Pilato & Lisi, 2006 and M. simulans Pilato et al., 2000. The new
species differs specifically from:
M. australis by: a different type of the oral cavity armature (harmsworthi-type in the new species and the echi-
nogenitus-type in M. australis), a lower pt of stylet supports (74.9–77.8 in the new species and ca. 83.7 in M. aus-
tralis), the egg surface covered by wrinkles (dots in M. australis), the process walls with small bubbles (smooth in
M. australis), a higher process height/base width ratio (1.51–1.61 in the new species and 0.95–1.05 in M. austra-
lis), a higher number of processes on the egg circumference (27–32 in the new species and 19–20 in M. australis),
a larger diameter of eggs without processes (85.1–94.5 in the new species and around 76.0 in M. australis) and with
processes (108.7–114.7 in the new species and around 96.0 in M. australis), narrower bases of egg processes (6.5–
9.0 in the new species and ca. 10.5 in M. australis).
M. coronatus by: a different type of the oral cavity armature (harmsworthi-type in the new species and the are-
olatus-type in M. coronatus), the egg surface covered by wrinkles (dots in M. coronatus), the process walls with
small bubbles (reticulated in M. coronatus), a higher process height/base width ratio (1.51–1.61 in the new species
and 0.88–0.96 in M. coronatus), a higher number of processes on the egg circumference (27–32 in the new species
and 11–18 in M. coronatus), a larger diameter of eggs without processes (85.1–94.5 in the new species and around
47.0–55.0 in M. coronatus) and with processes (108.7–114.7 in the new species and around 61.0–71.0 in M. coro-
natus)
M. patiens by: the presence of eyes, a larger body size (359–647 in the new species and 240–350 in M.
patiens), a lower pt of the buccal tube external width (12.0–13.2 in the new species and 17.9–21.1 in M. patiens), a
lower pt of the second macroplacoid length (7.6–8.9 in the new species and 11.9–13.8 in M. patiens), a lower pt of
the microplacoid length (5.5–7.4 in the new species and 10.4–12.9 in M. patiens), a lower pt of the macroplacoid
row length (34.4–40.8 in the new species and 42.7–48.9 in M. patiens), a lower pt of the placoid row length (42.3–
49.6 in the new species and 55.1–62.5 in M. patiens), the egg surface covered by wrinkles (dots in M. patiens), the
process walls with small bubbles (reticulated in M. patiens), a higher process height/base width ratio (1.51–1.61 in
the new species and ca. 0.65–1.05 in M. patiens), a higher number of processes on the egg circumference (27–32 in
the new species and 10–16 (usually 12) in M. patiens), a larger diameter of eggs with processes (108.7–114.7 in the
new species and 90.5–100.0 in M. patiens), narrower bases of egg processes (6.5–9.0 in the new species and 12.4–
19.0 in M. patiens).
M. pseudocoronatus by: having smooth cuticle (small dorsal and lateral tubercles present in M. pseudocorona-
tus), the absence of teeth on lunules IV, a lower pt of the buccal tube external width (12.0–13.2 in the new species

34 · Zootaxa 2781 © 2011 Magnolia Press KACZMAREK ET AL.

and 17.5–18.4 in M. pseudocoronatus), a lower pt of the second macroplacoid length (7.6–8.9 in the new species
and 11.7–13.6 in M. pseudocoronatus), a lower pt of the third macroplacoid length (10.2–12.5 in the new species
and 15.0–18.4 in M. pseudocoronatus), a lower pt of the microplacoid length (5.5–7.4 in the new species and 11.4–
11.5 in M. pseudocoronatus), a lower pt of the macroplacoid row length (34.4–40.8 in the new species and 47.3–
52.4 in M. pseudocoronatus), a lower pt of the placoid row length (42.3–49.6 in the new species and 60.0–65.1 in
M. pseudocoronatus), the egg surface covered by wrinkles (dots in M. pseudocoronatus), the walls of processes
with small bubbles (reticulated in M. pseudocoronatus), a higher process height/base width ratio (1.51–1.61 in the
new species and ca. 0.91–0.95 in M. pseudocoronatus), a higher number of processes on the egg circumference
(27–32 in the new species and 14 in M. pseudocoronatus), a larger diameter of eggs without processes (85.1–94.5
in the new species and around 50.1 in M. pseudocoronatus) and with processes (108.7–114.7 in the new species
and ca. 82.3 in M. pseudocoronatus), narrower bases of egg processes (6.5–9.0 in the new species and 11.5–13.9 in
M. pseudocoronatus).
M. radiatus by: the presence of eyes, a lower pt of the buccal tube external width (12.0–13.2 in the new species
and 17.5–20.1 in M. radiatus), a lower pt of the second macroplacoid length (7.6–8.9 in the new species and 13.5–
14.2 in M. radiatus), a lower pt of the third macroplacoid length (10.2–12.5 in the new species and 13.8–17.6 in M.
radiatus), a lower pt of the microplacoid length (5.5–7.4 in the new species and 10.9–11.3 in M. radiatus), a lower
pt of the macroplacoid row length (34.4–40.8 in the new species and 46.1–51.4 in M. radiatus), a lower pt of the
placoid row length (42.3–49.6 in the new species and 58.5–64.1 in M. radiatus), the egg surface covered by wrin-
kles (stripes forming a large radiate crown in M. radiatus), the process walls with small bubbles (reticulated in M.
radiatus), a higher number of processes on the egg circumference (27–32 in the new species and ca. 11 in M. radia-
tus)
M. rigidus by: a different type of the oral cavity armature (harmsworthi-type in the new species and the areola-
tus-type in M. rigidus), the presence of eyes, the absence of teeth on lunules on IV, a lower pt of the buccal tube
external width (12.0–13.2 in the new species and 17.5–20.1 in M. rigidus), a lower pt of the second macroplacoid
length (7.6–8.9 in the new species and 13.5–14.2 in M. rigidus), a lower pt of the third macroplacoid length (10.2–
12.5 in the new species and 13.8–17.6 in M. rigidus), a lower pt of the microplacoid length (5.5–7.4 in the new spe-
cies and 10.9–11.3 in M. rigidus), a lower pt of the macroplacoid row length (34.4–40.8 in the new species and
46.1–51.4 in M. rigidus), a lower pt of the placoid row length (42.3–49.6 in the new species and 58.5–64.1 in M.
rigidus), the egg surface covered by wrinkles (dots in M. rigidus), the walls of processes with small bubbles (retic-
ulated in M. rigidus), a higher process height/base width ratio (1.51–1.61 in the new species and ca. 1.05 in M. rigi-
dus), a higher number of processes on the egg circumference (27–32 in the new species and 12 in M. rigidus), a
larger diameter of eggs without processes (85.1–94.5 in the new species and ca. 64.0 in M. rigidus) and with pro-
cesses (108.7–114.7 in the new species and around 91.0 in M. rigidus), narrower bases of egg processes (6.5–9.0 in
the new species and 14.5–15.2 in M. rigidus).
M. simulans by: the absence of teeth on lunules IV, a larger body size (359–647 in the new species and 280–
360 in M. simulans), a lower pt of the buccal tube external width (12.0–13.2 in the new species and 17.6–21.1 in M.
simulans), a lower pt of the second macroplacoid length (7.6–8.9 in the new species and 11.2–12.8 in M. simulans),
a lower pt of the microplacoid length (5.5–7.4 in the new species and 9.6–11.7 in M. simulans), a lower pt of the
placoid row length (42.3–49.6 in the new species and 52.4–59.2 in M. simulans), the egg surface covered by wrin-
kles (dots in M. simulans), the walls of processes with small bubbles (reticulated in M. simulans), a higher process
height/base width ratio (1.51–1.61 in the new species and around 0.69–1.00 in M. simulans), a higher number of
processes on the egg circumference (27–32 in the new species and 11–13 in M. simulans), a larger diameter of eggs
without processes (85.1–94.5 in the new species and 43.0–82.0 in M. simulans) and with processes (108.7–114.7 in
the new species and 59.0–99.0 in M. simulans), narrower bases of egg processes (6.5–9.0 in new species and 11.0–
16.0 in M. simulans).

Key to the species of the harmsworthi group

1. Cuticle either with pores, or granulation, or stripes of pigmentation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

-. Cuticle smooth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
2(1). Pores in cuticle present, granulation and stripes of pigmentation absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
-. Pores in cuticle absent, granulation or stripes of pigmentation present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3(2). Egg shell smooth, processes in the shape of low cones (mammillate-like) without finger-like structures at the process base,
bubble-like structures at the top of processes present, 16–21 processes on the egg circumference . . . . . . . . . . . . . . . . . . . . . . . .

MACROBIOTUS BINIEKI SP. NOV. Zootaxa 2781 © 2011 Magnolia Press · 35

 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. krynauwi Dastych & Harris, 1995
-. Egg shell covered with dots, processes in the shape of high cones with finger-like structures at the process base, bubble-like
structures at the top of processes absent, 6–8 processes on the egg circumference . . . . . . . . . . . . . . M. liviae Ramazzotti, 1962
4(2). Nine bands of brown pigmentation on cuticle present* . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. baltatus McInnes, 1991
-. Cuticle without bands of pigmentation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5(4). Granulation on cuticle in the shape of large tubercles (1.5–2.0), present only on the caudal end of the body . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .M. arguei Pilato & Sperlinga, 1975
-. Granulation present on the entire dorsal side of the body. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6(5). Granulation on cuticle in the shape of small tubercles, eyes present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
-. Granulation on cuticle in the shape of fine dots; eyes absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
7(6). First band of teeth in the oral cavity armature and granulation on legs I–III absent, lunules IV smooth . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. perfidus Pilato and Lisi, 2009
-. First band of teeth in the oral cavity armature and granulation on legs I–III present, lunules IV dentate . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. pseudocoronatus Pilato et al., 2006
8(6). Areolae around the bases of egg processes absent, egg processes smooth, in the shape of spines with a wide base and a sharp
apex, processes smooth, 22–23 processes on the egg circumference . . . . . . . . . . . . . . . . . . . . . . . M. contii Pilato & Lisi, 2006
-. Areolae around the bases of egg processes present, egg processes in the shape of cones with a blunt apex, processes reticulated,
10–12 processes on the egg circumference . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. pseudonuragicus Pilato et al., 2004
9(1). Spurs on claws present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. armatus Pilato & Binda, 1996
-. Spurs on claws absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10(9). Egg processes situated very close (in contact) one to another . . . . . . . . . . . . . . . . . Macrobiotus neuquensis Rossi et al., 2009
-. Spaces between egg processes present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11(10). Areolation on eggs present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
-. Areolation on eggs absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
12(11). Each process inside a single areola . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .M. mauccii Pilato, 1974
-. A number of areolae present around each process . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13(12). Egg areolation poorly visible (areolae with thin walls), sometimes not fully developed, processes with a blunt apex . . . . . . 14
-. Egg areolation fully developed and well visible, processes with a sharp or indented apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
14(13). Eyes present, bubble-like structures at the top of processes absent, egg processes in the shape of cones, egg shell covered by
delicate stripes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. harmsworthi obscurus Dastych, 1985
-. Eyes absent, bubble-like structures at the top of processes present, egg processes in the shape of a bells (domes), egg shell cov-
ered by delicate dots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. peterseni Maucci, 1991
15(13). Body red, the oral cavity armature of the echinogenitus-type, lunules IV not thickened and without indentations, a crown of
dots around the bases of processes present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. ovostriatus Pilato & Patanè, 1998
-. Body transparent, the oral cavity armature of the harmsworthi-type, lunules IV thickened or with indentations, a crown of dots
around the bases of processes absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
16(15). Sixteen areolae around the base of each egg process . . . . . . . . . . . . . . . . . . . . . . . . . . . . .M. pseudoliviae Pilato & Binda, 1996
-. Six areolae around the base of each egg process . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
17(16). Process top divided into a number of points, more than 13 processes on the egg circumference . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. nuragicus Pilato & Sperlinga, 1975
-. Process top with 1–2 points, less than 13 processes on the egg circumference . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
18(17). Eyes and granulation on legs I present, lunules IV thickened, pt of stylet supports less than 76%, egg processes without a flex-
ible upper portion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. hieronimi Pilato & Claxton, 1988
-. Eyes and granulation on legs I absent, lunules IV with indentations, pt of stylet supports more than 77%, egg processes with a
flexible upper portion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .M. barbarae Kaczmarek et al., 2007
19(11). Egg processes hemispherical. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
-. Egg processes conical . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
20(19). Egg shell with reticular sculpture. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. montanus Murray, 1910
-. Egg shell without reticular sculpture. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .M. mottai Binda & Pilato, 1994
21(19). A well visible crown of swellings, dots or stripes around the bases of egg processes present . . . . . . . . . . . . . . . . . . . . . . . . . 22
-. A crown of swellings, dots or stripes around the bases of egg processes absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
22(21). Egg processes smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. australis Pilato & D'Urso, 1976
-. Egg processes reticulated or with small round bubbles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
23(22). A crown around the bases of egg processes composed of rectangular swellings, processes in the shape of spines with a wide
base, more than 25 processes on the egg circumference. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. binieki sp. nov.
-. A crown around the bases of egg processes composed of dots or stripes, processes in the shape of low cones, less than 20 pro-
cesses on the egg circumference . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
24(23). A crown around the bases of egg processes composed of radiate stripes forming a larger radiate crown, processes with a blunt,
indented apex. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. radiatus Pilato et al., 1991
-. A crown around the bases of egg processes composed of dots, processes with a sharp apex . . . . . . . . . . . . . . . . . . . . . . . . . . 25
25(24). Weakly visible granulation present only on legs IV, terminal portion of egg processes not flexible . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .M. rigidus Pilato & Lisi, 2006
-. Evident granulation present on legs I–IV, terminal portion of egg processes flexible . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
26(25). Eyes absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. patiens Pilato et al., 2000

36 · Zootaxa 2781 © 2011 Magnolia Press KACZMAREK ET AL.

-. Eyes present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
27(26). Additional teeth in the oral cavity absent, lunules IV indented . . . . . . . . . . . . . . . . . . . . . . . . . . M. simulans Pilato et al., 2000
-. Additional teeth in the oral cavity present, lunules IV smooth. . . . . . . . . . . . . . . . . . . . . . . . . . . . M. coronatus de Barros, 1942
28(21). Egg processes with bubble-like structures at the top of each process . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
-. Egg processes without bubble-like structures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
29(28). Finger-like structures around the bases of egg processes present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
-. Finger-like structures around the bases of egg processes absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
30(29). Lunules IV indented, more than 14 processes on the egg circumference . . . . . . . . . . . . . . . . . . . . . . . M. blocki Dastych, 1984
-. Lunules IV without indentation, less than 10 processes on the egg circumference . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
31(30). Eyes absent, oral cavity armature of the echinogenitus-type . . . . . . . . . . . . . . . . . . . . . . . . . M. tehuelchensis Rossi et al., 2009
-. Eyes present, oral cavity armature of the areolatus-type . . . . . . . . . . . . . . . . . . . M. szeptyckii Kaczmarek & Michalczyk, 2009
32(29). A few bubble-like structures at the top of each egg process, terminal filaments on processes present . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. stellaris du Bois-Reymond Marcus, 1944
-. Only one large bubble-like structure at the top of each egg process, terminal filaments on processes absent . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .M. reinhardti Michalczyk & Kaczmarek, 2003
33(28). Finger-like structures around the base of each egg process present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
-. Finger-like structures around the bases of egg processes absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
34(33). Eyes and the first band of teeth in the oral cavity present, lunules IV indented, egg process top smooth . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .M. erminiae Binda & Pilato, 1999
-. Eyes and the first band of teeth in the oral cavity absent, lunules IV smooth, egg process top reticulated . . . . . . . . . . . . . . . . 35
35(34). The pt ratio of stylet supports greater than 77%, lengths of all placoids more than 5.0 µm, pt of the macroplacoid row length
more than 50%, pt of the placoid row length more than 65% in adults . . . . . . . . . . . . . . . . M. snaresensis Horning et al., 1978
-. The pt ratio of stylet supports less than 76%, lengths of all placoids less than 3.0 µm, pt of the macroplacoid row length less
than 40%, pt of the placoid row lengthless than 50% in adults . . . . . . . . . . . . . . . . . . . . . . . . M. diguensis Pilato & Lisi, 2009
36(33). Eggs shell covered with radial striae, more than 15 processes on egg circumference . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
-. Egg shell smooth or covered with dots, less than 14 processes on egg circumference . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
37(36). Eyes absent, external claws IV less than 14.0 μm long, egg processes ending with one or a few very thin, flexible filaments . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. diffusus Binda & Pilato, 1987
-. Eyes present, external claws IV more than 22.0 μm long, egg processes without terminal filaments . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. barabanovi Tumanov, 2005
38(36). Two types of processes (spines and hemispheres) on egg present . . . . . . . . . . . . . . . . . . . . . . . . . . . M. wauensis Iharos, 1973a
-. Egg processes in the shape of cones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
39(38). Lunules IV smooth, egg shell smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. baltatus McInnes, 1991
-. Lunules IV indented, egg shell covered with dots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
40(39). Eyes present, egg processes in the shape of low cones (processes height/width ratio ca. 0.43–0.55) . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. harmsworthi harmsworthi Murray, 1907
-. Eyes absent, egg processes in the shape of high cones (processes height/width ratio ca. 1.25–1.40) . . M. altitudinalis Biserov,
1998
*Bands of pigmentation may become invisible after preparation.

Conclusion

We have found that a small survey of tardigrades from the Sofia City Province, Bulgaria reveals two species new to
the country and a species new to science. The new species, Macrobiotus binieki, belongs to the M. harmsworthi
group. Comparative analysis and compilation of a key to this group showed taxonomic detail (e.g. differences in
the oral cavity armature) indicating that this group requires further research, especially regarding its phylogenetic
nature. Further study of Bulgarian tardigrades is also required to elevate the knowledge of this regions fauna.

Acknowledgements

We are grateful to Peter Degma (Comenius University, Slovakia), Paul Bartels (Warren Wilson College, USA), an
anonymous referee and Sandra McInnes (British Antarctic Survey, UK) for their valuable comments on the manu-
script. This work was partially supported by funds from the Foundation for Polish Science (FNP) to ŁK.

MACROBIOTUS BINIEKI SP. NOV. Zootaxa 2781 © 2011 Magnolia Press · 37

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