On the diversity of abyssal Dondersiidae (Mollusca: Aplacophora) with the description of a new genus, six new species, and a review of the family
Author
Cobo, M. Carmen
0000-0002-8481-2086
Department of Zoology, Genetics and Physycal Antrhopology, University of Santiago de Compostela. Rúa Lope Gómez de Marzoa, s / n. Campus Vida. 15782 Santiago de Compostela, Spain. & Department of Biological Sciences and Alabama Museum of Natural History, University of Alabama. 300 Hackberry Lane, Tuscaloosa, AL 35487, USA. & kmkocot @ ua. edu; https: // orcid. org / 0000 - 0002 - 8673 - 2688
mariadelcarmen.cobo@usc.es
Author
Kocot, Kevin M.
0000-0002-8673-2688
Department of Biological Sciences and Alabama Museum of Natural History, University of Alabama. 300 Hackberry Lane, Tuscaloosa, AL 35487, USA. & kmkocot @ ua. edu; https: // orcid. org / 0000 - 0002 - 8673 - 2688
kmkocot@ua.edu
text
Zootaxa
2021
2021-02-18
4933
1
63
97
journal article
8067
10.11646/zootaxa.4933.1.3
1ec7cff3-1bd4-4038-87fc-8a6d87f4c7ef
1175-5326
4547986
303F97F8-463C-4A52-B5D7-28154E492493
Helluoherpia vieiralaneroi
sp. n.
(
Figure 6
,
Table 2
)
Type material.
Holotype
:
ZSM
Mol
20171268. (Zoologische Staatssammlung München).
Serial
sections (seven slides) and sclerites (one
SEM
stub, five slides).
Brazil
Basin
,
DIVA 3
Me
79/1 area 2, station 561 (
26º 34.78’S
,
035º 13.90’W
),
4484.7 to 4503 m
depth
.
Derivatio nominis.
Male genitive in honor of Dr. Vieira Lanero (University of Santiago de Compostela).
Diagnosis.
Small animal (<
2 mm
) with elongate body. Appearance slightly scaly. With a depression around the atrio-buccal cavity. With lanceolate, leaf-shaped scales as the main
type
of sclerites, pedunculate, lanceolate scales, and solid, curved acicular sclerites. With a single pedal fold. Common atrio-buccal cavity is very small and without papillae.
Helluoherpia
-
type
ventrolateral foregut glands. Radula with three denticles. Mantle cavity very small. With seminal vesicles.
Description.
Habitus
: Small animal (
1.35 mm
long,
0.2 mm
wide in the middle) with the anterior end rounded and somewhat wider than the rest of the body (
0.3 mm
). With a small ventral depression where the atrio-buccal opening is located (
Figure 6 A
). The posterior end was externally damaged, but good histological preparations were still obtained. The pedal groove is evident externally. Yellowish-white in 96 % ethanol. The imbricate arrangement of the sclerites is easily observed, although it does not have a very scaly appearance.
Mantle
: Thin epidermis (1.5 to 2.5 μm thick) without epidermal papillae. Thin cuticle (5 to 7.5 μm) with three
types
of sclerites arranged in a single layer: 1) Regular lanceolate, leaf-shaped scales (
Figure 6 B, C
) that cover the entire body (60 to 80.5 μm long, 20 to 12.5 μm wide). The other sclerites appear intermittently between them. 2) Irregular, pedunculate, lanceolate leaf-shaped scales (
Figure 6 D
) are the second most common
type
of sclerites (34 to 36 μm long, 10 to 12 μm wide). 3) Solid, acicular, sclerites (
Figure 6 E, D
) are present, concentrated mainly in the dorsal region. These are curved sclerites with a narrow proximal region and are somewhat flat (but rounded in cross-section) in the middle region, with a pointed end (60 to 65 μm long, 0.8 to 1.5 μm wide). These acicular sclerites are concentrated mainly in the dorsal region. They are the least abundant
type
. No sclerites specific to the pedal groove were found.
FIGURE 6.
Helluoherpia vieiralaneroi
sp. n.
A.
Habitus (holotype).
B.
Leaf-shaped scales.
C.
Regular lanceolate leaf-shaped scales.
D.
Irregular pedunculate lanceolate leaf-shaped scales.
E.
Solid acicular sclerites.
F.
Anatomical reconstruction of the anterior body based on manual reconstructions of the holotype.
G.
Anatomical reconstruction of the posterior body based on manual reconstructions of the holotype.
1-6.
Serial sections; location in the body is indicated in F and G.
H.
Reconstruction of the radula.
H’
Denticles in the position they were found.
H’’
Radular base.
Pedal groove and mantle cavity
: The pedal pit is very glandular (
Figure 6
F-2). It appears just posterior to the atrium and continues internally (25 μm long with an opening that is 10 μm in diameter, 10 to 32 μm wide, 5 to 10.5 μm high). The anterior pedal glands connect with the dorsal wall of the pedal pit. They are very voluminous ven-trally and reach the mantle cavity. The pit originates as a single, triangular pedal fold in the anterior region (7.5 μm wide 8 μm high), which flattens in the medial and posterior body regions (10.5 to 17.5 μm wide, 7.5 μm high).
The mantle cavity opening is ventro-posterior and very narrow (5 to 6 μm;
Figure 6
G-5). In the sections it looks like a folded duct as it is small (20 μm long, 27.5 to 30 μm wide, 32.5 μm high) and is virtually confined to the opening of the spawning duct (
Figure 6
G-5).
Digestive system
: The mouth, which is located at the posterior end of the atrium, continues as a foregut that is narrow and circular in cross-section (32.5 μm long, 5 to 15 μm in diameter;
Figure 6
F-2) and surrounded by a thin layer of longitudinal musculature (<2.5 μm). It also has a small dorsal pouch (12.5 μm long, 15 μm wide, 10 to 15 μm high).
Radula with three straight, identical narrow denticles per tooth (<1 to1.2 μm wide, fragments of 10 μm long) (
Figure 6
H-H´). Fragments of a straight radular base were also observed (2 to 8 μm anterior to posterior and 2 μm high) (
Figure 6
H-H´´). The fragmented tooth was observed in the radular sac. In this area, the ventrolateral foregut glands (
type
A) open ventrally in the foregut as a single duct (45 μm long). The ducts of the ventrolateral foregut glands are long and surrounded by a strong layer of longitudinal and circular musculature (approximately 5 μm thick). Most of the glandular cells discharge into the middle of the ducts, with the posterior region of the ducts free of glandular cells. The ventrolateral foregut glands are characteristic and were described for the first time for the
type
species of the genus
Helluoherpia
(
H. aegiri
) (
Handl & Büchinger 1996
) so they can be referred to as
Helluoherpia
-
type
. The midgut has a short dorsal caecum (10 μm long, 20 μm wide), just below where the esophagus ends. This caecum is circular in cross-section and similar in diameter to the foregut. The rectum emerges dorsally in the mantle cavity (5 to 5.5 μm in diameter) and its wall is made up of sparsely ciliated.
Nervous system and sense organs
: The cerebral ganglion (32.7 μm long, 62.5 to 82.5 μm wide, 32.5 to 70 μm high;
Figure 6
F-2) and most of the ganglia were easily observed. The pedal ganglia (5 μm long, 5 μm in diameter) are located on both sides of the pedal pit and are joined by a small (<2 μm wide) commissure. The supra-rectal com-missure is short and narrow (7.5 μm long, 5 to 6 μm thick). The atrium is very small (15 μm aperture and full length, 25 μm wide and 10 to 15 μm high) with no atrial papillae, but with a glandular wall (
Figure 6
F-1).
Gonopericardial system
: Well-formed gonads with the reproductive cells (lateral oocytes and spermatozoa in the central area) concentrated in their anterior region. The pericardium is long and narrow (217 μm long, 22.5 μm wide, 7 to 30 μm high;
Figure 6
G-3), with a short heart attached to its dorsal wall. The pericardioducts arise in the mid-posterior region of the pericardium; they form a large seminal vesicle (50 μm long, 25 to 30 μm wide and 30 to 55 μm high;
Figure 6
G-4) that is directed toward the posterior end, and although it has an unpaired origin, the posterior region is bi-lobed. The pericardioducts run parallel to the pericardium and the midgut, on both sides of the spawning duct as a pair of circular tubes (120 μm long and 7.5 to 12.5 μm in diameter;
Figure 6
G-3). The spawning duct is unpaired (175 μm long) in all its extension. The pericardioducts are centrally connected to its anterior region (52.5 μm wide, 30 to 45 μm high:
Figure 6 G
), which is narrower than the middle region (70 to 75 μm in diameter). The spawning duct narrows again in its posterior region (30 to 40 μm in diameter) and opens into the mantle cavity as a wide ciliated duct (opening 25 to 30 μm wide and high), which loses the surrounding musculature layer (2.5 to 4 μm). Without copulatory stylets.
Remarks.
The classification of
Helluoherpia vieiralaneroi
sp. n.
within
Dondersiidae
is determined by the
type
of mantle sclerites, the ventrolateral foregut glands (
type
A), the radula, and the absence of respiratory folds. The description of the radula of
H. vieiralaneroi
sp. n.
is based on fragments found in the radular sac and it is possible to affirm the presence of three denticles, which have an arrangement on the radula consistent with the genus
Helluoherpia
(
Handl & Büchinger 1996
)
. The inclusion of the new species in this genus is due to: 1) the sclerites (leaf-shaped scales and solid needles); 2) the radula; 3) the lack of a dorsoterminal sensory organ, copulatory stylets and respiratory folds; 4) the presence of a seminal vesicle, and 5) the peculiarities of the ventrolateral foregut glands (
type
A;
Helluoherpia
-
type
).
Helluoherpia vieiralaneroi
sp. n.
is considered a new species based on its geographical distribution and a combination of anatomical characters that set it apart from the only other known species of the genus,
Helluoherpia aegiri
. Externally it resembles
H. aegiri
although the specimen studied here is much smaller (
1.3 mm
) than the
holotype
of
H. aegiri
(
6 mm
;
Handl & Büchinger 1996
). The main type of leaf-shaped scales of the two species are similar in shape and size, but in the new species the solid acicular sclerites are curved and somehow flattened, while those of
H. aegiri
are straight (
Handl & Büchinger 1996
), and it also has fewer pedunculated leaf-shaped scales. The basic internal anatomical characteristics of
H. vieiralaneroi
sp. n.
and
H. aegiri
are similar but there are some important differences: a)
H. vieiralaneroi
sp. n.
has a very small atrium, with no atrial papillae, whereas in the atrium of
H. aegiri
, there are large globular papillae (
Handl & Büchinger 1996
); b) In
H. vieiralaneroi
sp. n.
the ducts of the ventrolateral foregut glands are shorter, and they run parallel to the foregut and then to the ventral region, and not to the dorsal region as in
H. aegiri
(
Handl & Büchinger 1996
)
; c)
H. vieiralaneroi
sp. n.
has a small midgut caecum that is absent in
H. aegiri
; d) In
H. vieiralaneroi
sp. n.
the seminal vesicles are exceptionally bulky and extended toward the posterior end, whereas in
H. aegiri
, they are smaller and extend toward the anterior region; e) The spawning duct in
H. vieiralaneroi
sp. n.
is much longer and bulkier than in
H. aegiri
(
Handl & Büchinger 1996
)
; f) The pericardium is markedly larger in
H. vieiralaneroi
sp. n.
than in
H. aegiri
, which has a very short pericardium (
Handl & Büchinger 1996
) and it also extends far posteriorly from where the pericardioducts fuse with it.