New records of tardigrades from Colombia with the description of Paramacrobiotus sagani sp. nov. and Doryphoribius rosanae sp. nov.
Author
Daza, Anisbeth
Author
Caicedo, Martín
Author
Lisi, Oscar
Author
Quiroga, Sigmer
text
Zootaxa
2017
2017-12-04
4362
1
29
50
journal article
31255
10.11646/zootaxa.4362.1.2
001e3a91-4ea0-4bb3-9ec8-9d3697554a85
1175-5326
1076122
F260162E-CB13-4B60-BF80-032E039D782F
Doryphoribius rosanae
sp.
nov.
(
Figs. 5–8
,
Table 4
)
Material examined:
Holotype
mounted in Hoyer’s medium,
1 paratype
in Hoyer’s medium, and
9 paratypes
in PVA from locality Bonda, lower basin of ManZanares river. Nineteen additional specimens were also found, 10 from the Puerto Mosquito locality, lower basin of the Gaira River, and 9 from the central basin of the Garupal River. All details about specimens are given in
Table 1
. The microhabitat, from which they were extracted, was mixed assemblages of bryophytes (
Dicranaceae
and
Racopilaceae
) and lichens (
Parmeliaceae
, Strigulaceae and
Thelotremataceae
) growing on tree trunks.
Holotype
found in a mixed assemblage of
Dicranaceae
and
Racopilaceae
species growing on tree.
Type
repositories:
The
holotype
and the
paratypes
are deposited in the Centro de Colecciones Biológicas de la Universidad del
Magdalena
(CBUMAG), Santa Marta,
Colombia
.
Holotype
CBUMAG:
TAR
:00009-3, and
10 paratypes
(
Table 1
).
Species diagnosis:
Colourless after mounting (not noticed before mounting). Eye-spots present. Dorso-lateral cuticle covered with a reticular design (
Fig. 5A, E–F
) that does not form tubercles at the point where the mesh crosses. Nine transversal rows of gibbosities on the dorso-lateral surface of the body; gibbosity configuration is IX:4-6-2-6-2-6-4-2-2 (
Fig. 5A
,
Fig. 6
). Rows II, IV and VI are aligned with legs I, II and III, respectively. Lateral gibbosities in lines I and VII misaligned, being more caudal, with respect to the dorsal gibbosities of the same row. Bucco-pharyngeal apparatus of
Doryphoribius
type
(
Fig. 5B
). Peribuccal lobes and papulae present. Oral armature with five to ten sparsely arranged large teeth in the region of the medio-dorsal ridge, and four to twelve sparsely arranged large teeth in the region of the medio-ventral ridge; in both cases the largest tooth is in the median position (
Fig. 7
). Stylet supports inserted at 67.6–72.8% of the buccal tube length. Pharynx with two elongated macroplacoids, the first longer, with slight central constriction. Microplacoids or septula absent. Claws of
Isohypsibius
type
, big, stout (
Fig. 5C–D
,
Fig. 8
); external (I–III) and posterior (IV) claws moderately bigger and with far larger base; free accessory points on main branches absent,
i.e.
very thin accessory points completely attached to the main branches present (
Fig. 8
, arrows). Narrow, smooth lunules present (
Fig. 5C
, arrows). Bars and other cuticular thickenings on legs absent.
Description of the
holotype
:
Body length 303 µm, colourless after mounting. Eye-spots present. Dorso-lateral cuticle covered with a reticular design (
Fig. 5A, E–F
), which appears quite rough and irregular with regards to the mesh shape and siZe, and the thickness of the delimiting ridges; where the mesh crosses it does not form tubercles. Nine rows of gibbosities on the dorso-lateral portion of the body (
Fig. 5A
,
Fig. 6
). The gibbosity configuration is IX:4-6-2-6-2-6-4-2-2. Rows II, IV and VI are aligned with legs I, II and III, respectively. The gibbosity arrangement is shown in
Fig. 6
. The first row shows two detached dorso-lateral gibbosities and two laterals in a more caudal position; rows from II to VII all show dorso-medial attached gibbosities plus dorso-lateral gibbosities, and lateral gibbosities in the rows II, IV and VI; the dorso-medial gibbosities tend to be flat and faint, especially those of rows III and V, and in several
paratypes
were difficult to observe. Lateral gibbosities in row VII misaligned, being more caudal, with respect to the dorsal gibbosities of the same row. Rows VIII and IX both have two dorso-lateral gibbosities moderately elongated and thus more protruding than the others, those of the last row almost pointed.
TABLE 4.
Measurements and
pt
values of selected morphological structures of the hotolotype and paratypes of
Doryphoribius rosanae
Sp.
nov.
| CHARACTER |
N |
RANGE |
MEAN |
SD |
Holotype |
| µm |
pt
|
µm
pt
|
µm |
pt
|
µm
pt
|
| Body length |
11 |
289 – |
434
–
|
340 |
48 |
303 |
| Buccopharyngeal tube |
| Buccal tube length |
10 |
33.3 – |
46.3
–
|
41.1
–
|
3.8 |
–
|
41.2
–
|
| Stylet support insertion point |
10 |
23.8 – |
31.4
67.6 –
|
72.8
|
28.9
70.3
|
2.4 |
1.5
|
30.0
72.8
|
| Buccal tube external width |
10 |
4.3 – |
6.4
11.1 –
|
14.5
|
5.2
12.6
|
0.7 |
1.2
|
5.0
12.1
|
| Buccal tube internal width |
9 |
2.3 – |
4.2
6.3 –
|
9.1
|
3.1
7.5
|
0.6 |
1.0
|
2.6
6.3
|
| Ventral lamina length |
8 |
17.2 – |
26.5
51.6 –
|
58.0
|
22.8
54.8
|
2.7 |
2.7
|
23.9
58.0
|
| Placoid lengths |
| Macroplacoid 1 |
11 |
5.3 – |
9.2
15.9 –
|
19.8
|
7.4
17.9
|
1.0 |
1.4
|
7.7
18.7
|
| Macroplacoid 2 |
11 |
3.4 – |
5.0
10.0 –
|
12.7
|
4.4
10.8
|
0.4 |
0.8
|
4.5
10.9
|
| Placoid row |
11 |
9.6 – |
15.1
28.4 –
|
34.3
|
12.6
31.0
|
1.5 |
1.8
|
14.1
34.3
|
| Claw 1 lengths |
| External base |
3 |
8.1 – |
8.6
20.6 –
|
20.6
|
8.4
20.6
|
0.3 |
0.1
|
?
?
|
| External primary branch |
3 |
12.1 – |
14.3
30.8 –
|
35.0
|
13.2
32.9
|
1.1 |
2.9
|
?
?
|
| External secondary branch |
3 |
10.1 – |
11.6
25.7 –
|
28.3
|
10.7
27.0
|
0.8 |
1.8
|
?
?
|
| Internal base |
8 |
5.5 – |
8.1
14.8 –
|
19.1
|
6.9
16.9
|
0.9 |
1.5
|
6.1
14.8
|
| Internal primary branch |
7 |
9.0 – |
14.0
27.0 –
|
32.4
|
11.9
29.4
|
1.9 |
2.5
|
11.1
27.0
|
| Internal secondary branch |
8 |
6.6 – |
10.9
20.0 –
|
25.7
|
9.1
22.4
|
1.4 |
1.8
|
9.5
23.1
|
| Claw 2 lengths |
| External base |
5 |
7.3 – |
10.5
20.9 –
|
22.9
|
8.9
22.1
|
1.3 |
0.9
|
?
?
|
| External primary branch |
4 |
12.0 – |
17.6
33.7 –
|
38.0
|
13.8
35.9
|
2.6 |
2.2
|
?
?
|
| External secondary branch |
6 |
8.7 – |
12.5
25.0 –
|
28.8
|
10.4
26.4
|
1.6 |
1.5
|
?
?
|
| Internal base |
5 |
5.4 – |
8.6
15.8 –
|
18.6
|
6.7
16.7
|
1.2 |
1.3
|
?
?
|
| Internal primary branch |
4 |
9.5 – |
15.2
28.4 –
|
32.9
|
12.7
31.2
|
2.5 |
2.4
|
?
?
|
| Internal secondary branch |
5 |
6.8 – |
9.5
19.0 –
|
20.6
|
8.2
20.0
|
1.0 |
0.7
|
?
?
|
| Claw 3 lengths |
| External base |
3 |
7.7 – |
10.3
23.0 –
|
25.1
|
9.0
24.1
|
1.3 |
1.5
|
?
?
|
| External primary branch |
3 |
11.4 – |
14.7
34.2 –
|
36.0
|
13.0
35.1
|
1.7 |
1.2
|
?
?
|
| External secondary branch |
4 |
8.9 – |
12.6
25.7 –
|
30.8
|
10.5
27.7
|
1.7 |
2.7
|
?
?
|
| Internal base |
4 |
5.6 – |
7.2
16.0 –
|
16.8
|
6.5
16.3
|
0.7 |
0.5
|
?
?
|
| Internal primary branch |
5 |
9.3 – |
13.4
26.5 –
|
30.9
|
11.4
28.8
|
1.8 |
1.9
|
?
?
|
| Internal secondary branch |
5 |
6.9 – |
9.3
20.6 –
|
22.1
|
8.1
21.4
|
0.9 |
0.6
|
?
?
|
| Claw 4 lengths |
| Anterior base |
4 |
6.4 – |
8.3
18.0 –
|
19.4
|
7.5
18.8
|
0.8 |
0.7
|
?
?
|
| Anterior primary branch |
4 |
10.5 – |
13.6
31.5 –
|
32.9
|
12.3
32.1
|
1.5 |
0.7
|
?
?
|
| Anterior secondary branch |
4 |
7.5 – |
10.3
22.4 –
|
24.1
|
9.0
23.4
|
1.2 |
0.9
|
?
?
|
| Posterior base |
5 |
7.6 – |
10.5
22.6 –
|
24.7
|
9.2
23.2
|
1.1 |
1.0
|
?
?
|
| Posterior primary branch |
4 |
10.7 – |
15.2
32.0 –
|
35.9
|
13.3
34.1
|
2.0 |
1.9
|
?
?
|
| Posterior secondary branch |
5 |
9.1 – |
11.9
26.7 –
|
28.0
|
10.7
27.2
|
1.1 |
0.5
|
?? |
FIGURE 5.
Doryphoribius rosanae
sp.
nov.
A, habitus (holotype). B, buccal apparatus (holotype). C, claws leg II (paratype CBUMAG:TAR:00225-7); the arrows indicate the lunules. D, claws leg IV (paratype CBUMAG:TAR:00227-9). E–F, dorsal cuticle surface with detail of some gibbosities (holotype). F: DIC, all others: PCM.
FIGURE 6.
Doryphoribius rosanae
sp.
nov.
Schematic depiction of the cuticular gibbosity configuration. Roman numbers indicate the transverse rows of gibbosities.
Bucco-pharyngeal apparatus of
Doryphoribius
type
, with ventral lamina long more than half the buccal tube (58.0% of the tube length) (
Fig. 5B
). Peribuccal lobes and papulae are present. Oral armature with two dorsolateral and two ventro-lateral ridges, both with sparsely arranged large teeth in the region of a median ridge: ten medio-dorsal teeth, and twelve medio-ventral teeth, with the largest tooth in the median position (
Fig. 7
). Buccal tube 41.2 µm long and externally 5.0 µm wide (
pt
=
12.1
). Pharyngeal bulb with apophyses and two elongated macroplacoids; microplacoids and septula absent. First macroplacoid 7.7 µm long (
pt
=
18.7
), with slight central constriction; second macroplacoid 4.5 µm long (
pt
=
10.9
), with subterminal constriction; entire placoid row, 14.1 µm long (
pt
=
34.3
). Claws of
Isohypsibius
type
, big and stout, with relatively long branches (
Fig. 5C–D
,
Fig. 8
); the internal or anterior claws moderately smaller and with smaller bases than the external or posterior claws, which have more elongated branches and wider bases. Primary branches with thin accessory points completely attached to the main branches, so that “free” accessory points are lacking (
Fig. 8
, arrows). Internal claws on leg I with primary branch 11.1 µm long (
pt
=
27.0
), secondary branch 9.5 µm long (
pt
=
23.1
), and base 6.1 µm long (
pt
=
14.8
). The orientation of all the other claws were unsuitable for measurement. Narrow, smooth lunules present under all claws (
Fig. 5C
, arrows), more developed under the external (I–III) and especially the posterior (IV) claws. Bars and other cuticular thickenings on legs absent.
The measurements of selected morphological structures of the
holotype
and the ranges within the population, are given in
Table 4
.
Taxonomic remarks:
The
paratypes
were similar to the
holotype
in both qualitative and metric characters, only the number of teeth in the buccal armature has shown variation, but this might also depend on the different orientation of the specimens, which could obscure some teeth; to give the range of variability, we were able to count from four to ten teeth in dorsal view, and five to twelve teeth in ventral view.
Etymology.
This species is dedicated to the biologist Rosana Londoño, colleague and friend of the research group, who has worked with us for many years on tardigrade taxonomy.
FIGURE 7.
Oral armature of
Doryphoribius rosanae
sp.
nov.
(holotype). A, the dorsal portion is better focused; the arrow indicates some dorsal teeth. B, the dorso-lateral portion is focused; arrows “a” indicate two dorsal teeth; arrow “b” indicates some dorso-lateral teeth corresponding to the lateral ridge. C, the ventral portion is focused, with several teeth visible. D, Image obtained by merging several photos with different focuses, so that both dorsal and ventral teeth are visible. Photos taken under PCM.
FIGURE 8.
Detail of the attached accessory point structure of
Doryphoribius rosanae
sp.
nov.
, claws of leg IV (paratype CBUMAG:TAR:00225-7); the arrows indicate the “accessory points” attached to the main branches. Photos taken under PCM.
Differential diagnosis.
The presence of cuticular gibbosities and two macroplacoids in the pharynx places
Doryphoribius rosanae
sp.
nov.
into the
evelinae
group (MichalcZyk & KacZmarek 2010). The reticular design on the dorso-lateral cuticle and nine rows of gibbosities is most similar to:
D
.
barbarae
Beasley & Miller, 2012
,
D
.
dawkinsi
MichalcZyk & KacZmarek, 2010
,
D
.
huangguoshuensis
Wang, Wang & Li, 2007
,
D
.
maasaimarensis
Fontoura,
Lisi & Pilato, 2013
,
D
.
niedbalai
Zawierucha,
MichalcZyk & KacZmarek, 2012
,
D
.
zyxiglobus
(
Horning, Schuster & Grigarick, 1978
)
, and
D
.
mcinnesae
Meng, Sun & Li, 2014
.
Doryphoribius rosanae
sp.
nov.
differs from all these species in having a unique gibbosity configuration (IX:4-6-2-6-2-6-4-2-2), and in lacking cuticular tubercles at the crossing of the cuticular reticular sculpture (this character though not described for
D. niedbalai
was clearly visible in
Fig. 5
of the species description,
Zawierucha
et al.
2012
).
Doryphoribius rosanae
sp.
nov.
differs from
D. barbarae
by having a larger body siZe (221.1–239.6 µm in
D. barbarae
, 289–434 µm in
D. rosanae
sp. nov.
), different number and arrangement of gibbosities (IX:2-4-4-4-4-6- 4-4-
2 in
D. barbarae
, IX:4-6-2-6-2-6-4-2-
2 in
D
.
rosanae
sp. nov.
), presence of eye-spots, and lunules.
Doryphoribius rosanae
sp.
nov.
differs from
D. dawkinsi
, by having a larger body siZe (165.0–266.2 µm in
D. dawkinsi
, 289–434 µm in
D
.
rosanae
sp. nov.
), different number and arrangement of gibbosities (IX:6-6-4-6-4-6-4- 4-2+2[L IV] in
D. dawkinsi
, IX:4-6-2-6-2-6-4-2-
2 in
D
.
rosanae
sp. nov.
), presence of eye-spots, and claws with lunules.
Doryphoribius rosanae
sp.
nov.
differs from
D. huangguoshuensis
by having a different number and arrangement of gibbosities (IX:4-4-4-4-4-4-4-2-
2 in
D. huangguoshuensis
, IX:4-6-2-6-2-6-4-2-
2 in
D
.
rosanae
sp. nov.
), and a wider buccal tube (
pt
8.3–8.8
in
D. huangguoshuensis
,
11.1–14.5
in
D
.
rosanae
sp. nov.
).
Doryphoribius rosanae
sp.
nov.
differs from
D. maasaimarensis
by having a larger body siZe (175–288 µm in
D. maasaimarensis
, 289–434 µm in
D
.
rosanae
sp. nov.
), different number and arrangement of gibbosities (IX:4-6- 4-6-4-6-4-4-
2 in
D. maasaimarensis
, IX:4-6-2-6-2-6-4-2-
2 in
D
.
rosanae
sp. nov.
), a wider buccal tube (
pt
8.7–9.4
in
D. maasaimarensis
,
11.1–14.5
in
D. rosanae
sp. nov.
), and the absence of ornamentation on legs.
Doryphoribius rosanae
sp.
nov.
differs from
D. niedbalai
by having a different number and arrangement of gibbosities (IX:4-6-5-6-5-6-4-2-
2 in
D. niedbalai
, IX:4-6-2-6-2-6-4-2-
2 in
D
.
rosanae
sp. nov.
), by the presence of eye-spots, and absence of free accessory points.
Doryphoribius rosanae
sp.
nov.
differs from
D. zyxiglobus
in the gibbosity configuration (IX:4-6-4-6-4-6-4-4-
2 in
D. zyxiglobus
, IX:4-6-2-6-2-6-4-2-
2 in
D
.
rosanae
sp. nov.
), a wider buccal tube (
pt
9.2–11.8
in
D. zyxiglobus
,
11.1–14.5
in
D. rosanae
sp. nov.
), larger macroplacoids (
pt
12.6–14.7
in first macroplacoid and
8.4–9.9
in
second macroplacoid in
D. zyxiglobus
,
15.9–19.8
and
10.0–12.7
, respectively, in
D. rosanae
sp. nov.
), and in having more robust claws (e.g.
pt
claw IV anterior, base
13.0–15.9
, primary branch
26.6–29.8
and secondary branch
19.4–21.9
in
D. zyxiglobus
, 18.0–19.4
,
31.5–32.9
and
22.4–24.1
, respectively, in
D. rosanae
sp. nov.
) with smaller, attached, accessory points (larger accessory points, with free tips in
D. zyxiglobus
).
According to the original description of
D. mcinnesae
,
D
.
rosanae
sp. nov.
would differ in having a different number of rows of gibbosities, and of gibbosities themselves (X:2-3-1-3-1-3-1-2-2-
2 in
D. mcinnesae
, IX:4-6-2-6- 2-6-4-2-
2 in
D
.
rosanae
sp. nov.
); however, we think
D. mcinnesae
can also be considered has having nine rows of gibbosities (see discussion), the formula for which would be IX:2-3-1-3-1-3-3-2-2, therefore still different from the new species. Additionally,
D. rosanae
sp. nov.
has lunules, which are lacking in
D. mcinnesae
.