Tardigrades from Peru (South America), with descriptions of three new species of Parachela
Author
Kaczmarek, Łukasz
Author
Cytan, Joanna
Author
Zawierucha, Krzysztof
Author
Diduszko, Dawid
Author
Michalczyk, Łukasz
text
Zootaxa
2014
3790
2
357
379
journal article
45952
10.11646/zootaxa.3790.2.5
7895e366-3da3-46a2-9d68-58acb2cf3f89
1175-5326
231292
564A86FD-557A-43CA-B015-6BA767E281F9
Paramacrobiotus intii
sp. nov.
(
Figs 18–32
;
Tables 4–5
)
Localities and number of specimens:
I (125+42, 20+5 SEM). Material examined:
Holotype
(slide PE2001/82) and 167
paratypes
(124 animals and
42 eggs
) (slides: PE2001/*, where the asterisk can be substituted by any of the following numbers: 4, 5, 6, 7, 8, 10, 12, 15, 17, 18, 19, 24, 31, 33, 35, 36, 37, 38, 40, 41, 42, 43, 44, 45, 46, 47, 48, 49, 50, 51, 52, 55, 56, 57, 59, 60, 61, 62, 64, 65, 71, 73, 74, 75, 76, 79, 80, 81, 82, 83).
Description
(measurements and statistics in
Tables 4–5
).
Animals:
Body colour transparent/white, eyes present in 87 of 125 (70%) fixed specimens (
Figs 18–19
). Except for all legs, where well visible granulation is present (
Figs 24–25
), cuticle is smooth,
i.e.
without gibbosities, papillae, pores, spines or sculpturing.
Bucco-pharyngeal apparatus of the
Macrobiotus
type
, with the ventral lamina and ten peribuccal lamellae (
Fig. 20–23
). Mouth antero-ventral. The oral cavity armature composed of two posterior bands of teeth (similar on dorsal and ventral side) (
Fig. 21–23
). The first/anterior band of teeth is absent or not visible under PCM. The second band of teeth, comprising small granules arranged in a few irregular rows that run around the oral cavity wall (PCM), is positioned at the rear of the oral cavity, between the ring fold and the third band of teeth. The third band of teeth is positioned just before the buccal tube opening and composed of large transverse ridges (PCM). In this band, three ventral and three dorsal teeth are present (two lateral and one median). The median tooth is sometimes divided into a few smaller teeth. Pharyngeal bulb spherical, with triangular apophyses and three rodshaped macroplacoids. Macroplacoid length sequence 2<1<3 (
Fig. 20
). The first macroplacoid without constrictions but distinctly narrower anteriorly. The second macroplacoid of a uniform width and without constrictions. The third macroplacoid with a sub-terminal constriction. Microplacoid absent.
Claws of the
hufelandi
type
, stout (
Figs 24–25
). Primary branches with distinct accessory points. Lunules under claws I–III smooth (
Fig. 24
) and distinctly dentate on legs IV (
Fig. 25
). Thin cuticular bars under claws I–III present. Other cuticular thickenings on legs absent.
Eggs:
Laid freely, white, spherical and ornamented (
Figs 19
,
26–32
). Processes in the shape of blunt or slightly sharpened cones (
Figs 19
,
26–27, 31–32
). Labyrinthine layer between process walls is visible under PCM as a clear reticular pattern (
Fig. 28
). Egg shell areolated with a single ring of ten areolae around each process (
Figs 29–30
). Internal surface of areolae smooth and without pores (PCM and SEM).
Etymology.
The new species is named after Inti, the ancient Incan Sun god.
Type
locality.
Peru
:
13°25'S
;
71°51'W
,
ca.
3,000 m
asl, Cusco Region, Pisac near Cusco, mixed mosses and lichens from rock, date:
28.10.2010
, coll. Dawid Diduszko.
Type
depositories.
Holotype
(slide slide PE2001/82) and 148
paratypes
(slides: PE2001/*, where the asterisk can be substituted by any of the following numbers: 4, 5, 6, 7, 8, 10, 12, 15, 17, 18, 19, 24, 31, 33, 35, 36, 37, 38, 40, 41, 42, 43, 44, 45, 46, 47, 48, 49, 50, 51, 52, 55, 56, 57, 59, 60, 61, 62, 64, 65, 71, 73, 74, 75, 76, 79, 80, 81, 82, 83) are deposited at the Department of Animal Taxonomy and Ecology, Institute of Environmental Biology, A. Mickiewicz University in Poznań, Umultowska 89, 61-614 Poznań (
Poland
), 10
paratypes
(PE2001/41, PE2001/ 47, PE2001/83) are deposited at the Natural History Museum, University of Copenhagen Universitetsparken 15, DK-2100 Copenhagen,
Denmark
and 9
paratypes
(PE2001/40, PE2001/49, PE2001/76) are deposited at the Department of Entomology Institute of Zoology Jagiellonian University, Gronostajowa 9, 30-387, Kraków,
Poland
.
Differential diagnosis.
By the shape of egg processes and by the lack of a microplacoid,
Paramacrobiotus intii
sp. nov.
is most similar to the species of the
areolatus
and
huziori
groups within the genus
Paramacrobiotus
,
i.e.
P. areolatus
(
Murray, 1907
)
,
P. centesimus
(Pilato, 2000)
,
P. crenatus
(
Maucci, 1991
)
,
P. derkai
(
Degma, Michalczyk & Kaczmarek, 2008
)
,
P. huziori
(
Michalczyk & Kaczmarek, 2006b
),
P. klymenki
Pilato, Kiosya, Lisi & Sabella, 2012
,
P. tonollii
(
Ramazzotti, 1956
)
, and
P. walteri
(
Biserov, 1997/8
), but it differs from all of them by a different
type
of the oral cavity armature (
i.e.
the presence of only two posterior bands of teeth, teeth of the second row in the shape of granules) in the new species
vs.
the
areolatus
oral cavity
type
(three bands of teeth present, teeth of the second row in the shape of small ridges parallel to the main axis of the buccal tube). Moreover,
Paramacrobiotus intii
sp. nov.
also differs from the abovementioned species:
FIGURES 18–19.
Paramacrobiotus intii
sp. nov.
—habitus
:
18
—adult (latero-ventral view, holotype);
19
—hatching juvenile (latero-ventral view, paratype). Both PCM.
FIGURES 20–25.
Paramacrobiotus intii
sp. nov.
—20
—buccal apparatus (dorso-ventral projection, holotype);
21-23
—oral cavity armature
(21
—ventral teeth of the second and the third band,
22
—dorsal teeth of the second band,
23
—dorsal teeth of the third band, scale on 22–23 same as on 21, holotype);
24
—claws III (paratype);
25
—claws IV (paratype). All PCM.
FIGURES 26–32.
Paramacrobiotus intii
sp. nov.
—egg
:
26
—embryonate egg (mid-section, PCM);
27
—chorion (SEM);
28
—labyrinthine layer between process walls (PCM);
29
—ten areolae around a process (PCM);
30
—areolation (SEM);
31–32
—processes (mid-sections, PCM).
TABLE 4.
Measurements [in µm] and
pt
values of selected morphological structures of
Paramacrobiotus intii
sp. nov.
mounted in Hoyer’s medium (N—number of specimens/structures measured, RANGE refers to the smallest and the largest structure among all measured specimens; SD—standard deviation,?—trait oriented unsuitably for measurement).
| CHARACTER |
N |
RANGE |
MEAN |
SD |
Holotype |
| µm pt |
µm
pt
|
µm
pt
|
µm
pt
|
| Body length |
15 |
346 – 575
819 – 1144
|
481
989
|
62
89
|
456
983
|
| Buccal tube |
| Length |
15 |
39.5 – 55.9
–
|
48.6
–
|
4.3
–
|
46.4
–
|
| Stylet support insertion point |
15 |
30.9 – 44.5
77.5 – 81.4
|
38.6
79.4
|
3.6
1.0
|
37.3
80.4
|
| External width |
15 |
5.5 – 9.1
13.9 – 16.9
|
7.6
15.5
|
0.9
0.9
|
7.2
15.5
|
| Internal width |
15 |
3.7 – 6.6
9.4 – 12.2
|
5.4
11.0
|
0.7
0.9
|
4.7
10.1
|
| Ventral lamina length |
5 |
24.2 – 33.5
56.6 – 65.7
|
28.7
61.5
|
3.8
3.7
|
29.0
62.5
|
| Placoid lengths |
| Macroplacoid 1 |
15 |
5.6 – 9.3
14.0 – 18.2
|
7.8
15.9
|
1.2
1.3
|
6.5
14.0
|
| Macroplacoid 2 |
15 |
4.5 – 7.2
11.2 – 13.4
|
5.9
12.1
|
0.8
0.7
|
5.5
11.9
|
| Macroplacoid 3 |
15 |
5.8 – 10.3
14.7 – 19.0
|
8.5
17.5
|
1.2
1.2
|
7.8
16.8
|
| Macroplacoid row |
15 |
16.8 – 27.3
42.5 – 53.4
|
23.8
48.7
|
3.1
2.7
|
22.4
48.3
|
| Claw 1 lengths |
| External primary branch |
8 |
10.0 – 14.0
24.2 – 27.8
|
12.4
25.7
|
1.4
1.1
|
12.1
26.1
|
| External secondary branch |
7 |
7.4 – 11.6
16.3 – 21.8
|
9.2
19.1
|
1.4
2.5
|
?
?
|
| Internal primary branch |
8 |
9.6 – 13.1
23.3 – 25.4
|
12.1
24.3
|
1.1
0.8
|
11.8
25.4
|
| Internal secondary branch |
8 |
8.1 – 11.4
18.2 – 20.9
|
9.9
19.8
|
1.1
1.0
|
8.7
18.8
|
| Claw 2 lengths |
| External primary branch |
8 |
11.1 – 14.6
25.0 – 29.2
|
13.3
27.8
|
1.2
1.4
|
13.4
28.9
|
| External secondary branch |
7 |
7.3 – 12.1
18.5 – 24.7
|
10.0
20.9
|
1.6
2.2
|
8.9
19.2
|
| Internal primary branch |
8 |
11.2 – 15.0
24.7 – 30.0
|
13.0
26.8
|
1.4
2.0
|
?
?
|
| Internal secondary branch |
8 |
9.1 – 12.8
21.2 – 26.3
|
11.2
23.1
|
1.4
1.9
|
?
?
|
| Claw 3 lengths |
| External primary branch |
8 |
10.5 – 16.0
24.4 – 29.7
|
13.3
27.6
|
1.8
1.6
|
13.5
29.1
|
| External secondary branch |
8 |
8.6 – 11.8
18.1 – 23.2
|
10.0
20.7
|
1.2
1.8
|
8.6
18.5
|
| Internal primary branch |
8 |
9.7 – 14.5
24.1 – 30.3
|
12.6
27.0
|
1.7
2.3
|
13.0
28.0
|
| Internal secondary branch |
7 |
8.7 – 11.6
20.6 – 24.6
|
10.4
22.2
|
1.2
1.6
|
?
?
|
| Claw 4 lengths |
| Anterior primary branch |
10 |
11.1 – 16.0
25.8 – 33.4
|
14.3
29.1
|
1.4
2.7
|
14.6
31.5
|
| Anterior secondary branch |
10 |
7.6 – 11.9
19.2 – 24.0
|
10.8
22.0
|
1.3
1.8
|
10.3
22.2
|
| Posterior primary branch |
12 |
10.7 – 16.6
26.1 – 34.7
|
14.7
30.2
|
1.7
2.5
|
15.0
32.3
|
| Posterior secondary branch |
12 |
8.4 – 12.6
18.2 – 26.8
|
10.7
22.0
|
1.0
2.4
|
10.4 22.4 |
Paramacrobiotus areolatus
, with a global distribution (
McInnes 1994
), by: the shape of egg process (processes with blunt or slightly sharpened apices in the new species
vs.
processes with clearly sharp apices in
P. areolatus
), a different process height/base width ratio (processes wider than longer in the new species
vs.
processes longer than wider in
P. areolatus
), and by slightly wider egg processes (22.0–34.0 Μm in the new species
vs.
19.0–22.0 Μm in
P. areolatus
).
Paramacrobiotus centesimus
, known only from
Brazil
(Pilato 2000), by: the presence of dentate lunules on claws IV, a different shape of apices of egg processes (blunt or slightly sharpened apices in the new species
vs.
sharp or indented apices in
P. centesimus
), slightly higher egg processes (15.4–24.4 Μm in the new species
vs.
7.0–11.0 Μm in
P. centesimus
), and by a slightly lower number of egg processes on the egg circumference (
9–10 in
the new species
vs.
11–12 in
P. centesimus
).
Paramacrobiotus crenatus
, known from
Greenland
and
Russia
(
Kaczmarek
et al.
2005
), by: a different shape of apices of egg processes (blunt or slightly sharpened apices in the new species
vs.
clearly sharp apices in
P. crenatus
), a different process height/base width ratio (processes wider than longer in the new species
vs.
processes longer than wider in
P. crenatus
), and by shorter egg processes (15.4–24.4 Μm in the new species
vs.
40.0–44.0 Μm in
P. crenatus
).
TABLE 5.
Measurements [in µm] of selected morphological structures of eggs of
Paramacrobiotus intii
sp. nov.
mounted in Hoyer’s medium (N—number of specimens/structures measured, RANGE refers to the smallest and the largest structure among all measured specimens; SD—standard deviation).
| CHARACTER |
N |
RANGE |
MEAN SD |
| Diameter of egg without processes |
13 |
75.3 – 113.5 |
89.5 10.2 |
| Diameter of egg with processes |
13 |
78.8 – 137.3 |
119.6 15.0 |
| Process height |
39 |
15.4 – 24.4 |
20.1 2.0 |
| Process base width |
39 |
22.0 – 34.0 |
27.3 3.1 |
| Process base/height ratio |
39 |
100% – 160% |
137% 20% |
| Distance between processes |
39 |
2.2 – 30.0 |
4.5 4.3 |
| Number of processes on the egg circumference |
13 |
9 – 10 |
9.7 0.5 |
Paramacrobiotus derkai
, known only from
Colombia
(
Degma
et al
. 2008
), by: less protruding accessory points, the presence of dentate lunules on claws IV, a different shape of apices of egg processes (blunt or slightly sharpened apices in the new species
vs.
clearly sharp and flexible tips in
P. de r k a i
), a different
type
of egg areolation (the
areolatus
type
(a circle of large and unbranched areolae separates adjacent conical egg processes, the space between neighbouring areolae usually smaller than their width) in the new species
vs.
the
huziori
type
(small and branched areolae separate adjacent egg processes, the space between neighbouring areolae usually broader than their width) in
P. derkai
), and by a slightly lower number of egg processes on the egg circumference (
9–10 in
the new species
vs.
12–16 in
P. d e r k ai
).
Paramacrobiotus huziori
known only from
Costa Rica
(
Michalczyk & Kaczmarek 2006b
, Kaczmarek
et al.
2014) by: a different shape of apices of egg processes (blunt or slightly sharpened apices in the new species
vs.
sharp and elongated apices covered by small tubercles in
P. h uz i or i
), and by a different
type
of egg areolation (the
areolatus
type
(a circle of large and unbranched areolae separates adjacent conical egg processes, the space between neighbouring areolae usually smaller than their width) in the new species
vs.
the
huziori
type
(small and branched areolae separate adjacent egg processes, the space between neighbouring areolae usually broader than their width) in
P. hu z i o r i
.
Paramacrobiotus klymenki
, known only from
Belarus
(
Pilato
et al
. 2012
), by: the presence of dentate lunules on claws IV, a different shape of egg processes (blunt or slightly sharpened apices with fully reticulated walls in the new species
vs.
sharp apices and with a small upper portion without wall reticulation in
P. klymenki
), and by slightly wider bases of egg processes (22.0–34.0 Μm in the new species
vs.
16.4–18.2 Μm in
P. klymenki
).
Paramacrobiotus tonollii
, with a mainly North American distribution (
McInnes 1994
,
Meyer 2013
), by: the absence of pores in the cuticle, the presence of dentate lunules on claws IV, the
type
of egg shell sculpture (areolation of the
areolatus
type
(a circle of large areolae around each process) in the new species
vs.
six ribs present at the bases of each process in
P. tonollii
), and by shorter egg processes (15.4–24.4 Μm in the new species
vs.
32.0–35.0 Μm in
P. tonollii
).
Paramacrobiotus walteri
, known only from
Russia
(
Biserov, 1997/8
) by: a different shape of apices of egg processes (blunt or slightly sharpened apices in the new species
vs.
clearly sharp and flexible bi- tri- or multifurcated apices in
P. walteri
).