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
Author
Kaczmarek, Łukasz
Author
Gołdyn, Bartłomiej
Author
Prokop, Zofia M.
Author
Michalczyk, Łukasz
text
Zootaxa
2011
2781
29
39
journal article
10.5281/zenodo.206999
f488c32f-c9a5-4c9c-9326-75a0ec9e287b
1175-5326
206999
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 continuous 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 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.
FIGURES 1–3.
Macrobiotus binieki
sp. nov.
adult: 1. habitus (ventral view, holotype), 2. buccal apparatus (ventral view, holotype), 3. claws III.
FIGURES 4–8.
Macrobiotus binieki
sp. nov.
egg: 4. mid-section, 5–6. egg surface (at two different focusing levels), 7–8. processes. Scale on 6 same as on 5 and scale on 8 same as on 7.
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 apophyses 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. Processes 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 (which could also be interpret as very small finger-like structures) (
Figs 5–6
). Processes are distributed on the surface 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 Animal 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
echinogenitus
-
type
in
M. australis
), a lower
pt
of stylet supports (
74.9–77.8
in
the new species and
ca
.
83.7
in
M. australis
), 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. australis
), 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
areolatus
-
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. coronatus
)
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. pseudocoronatus
), the absence of teeth on lunules IV, a lower
pt
of the buccal tube external width (
12.0–
13.2
in the new species 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 wrinkles (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. radiatus
)
M. rigidus
by: a different
type
of the oral cavity armature (
harmsworthi
-
type
in the new species and the
areolatus
-
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 species 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 (reticulated in
M. rigidus
), a higher process height/base width ratio (
1.51–1.61 in
the new species and ca.
1.05 in
M. rigidus
), 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 processes (
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 wrinkles (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
).