Some Marine Rotifera from Réunion Island, with a
Description of a New Species of Lindia
Harring and Myers, 1924 and one of Synchaeta Ehrenberg, 1832
Willem H. De Smet
Department of Biology, Section Polar
Ecology, Limnology and Palaeobiology,
University of Antwerp, Campus Middelheim, Groenenborgerlaan 171, B-2020 Antwerpen, Belgium. E-mail: willem.desmet@ua.ac.be
(Accepted June 3,
2005)
*To whom correspondence and
reprint requests should be addressed.
E-mail:
willem.desmet@ua.ac.be
Abstract Willem H. De Smet(2005) Some Marine Rotifera from
Key
words: Rotifera, Marine, Lindia elsae, Synchaeta squamadigitata, New species.
----------------------------------------------------------------------------------------------------------------
Introduction
To the present, the marine rotifer
fauna of the Southern Hemisphere has largely been neglected, with scarce
information being mostly restricted to occasional samplings. For the Ethiopian region Brownell (1988)
described Synchaeta hutchingsi from
the S.E. Atlantic off
----------------------------------------------------------------------------------------------------------------
MATERIALS AND
METHODS
Réunion (21°S, 55.30°E) is one of
the Mascarene Is. lying in the Indian Ocean, ~
Fig. 1.
Map showing
The island was visited on
the occasion of expeditions to the Subantarctic, organized by the
"Institut Polaire Français, Paul Emile Victor", and 2 marine samples
were taken.
Sample 1. Aufwuchs collected among a stand of the brown alga Padina pavonia (L.) Gaillon, from the infralittoral fringe of
Grande Anse at Grand Bois, S. Réunion on 24 Dec. 1998. Fixation was done on the spot with
formalin.
Sample 2. A vertical
plankton haul of
Animals were examined and
drawn with a Leitz Orthoplan microscope equipped with a camera lucida. Trophi were isolated by dissolving the
soft body parts in dilute NaOCl, and were prepared for light and scanning
electron microscopy (SEM) following the procedure of De Smet (1998). SEM was performed using a Philips SEM
515 microscope operated at 20 kV.
----------------------------------------------------------------------------------------------------------------
RESULTS
In tota l2 taxa were identified
(Table 1). Both samples contained
bdelloids (deformed beyond identification due to contraction) and
monogononts. Monogononts dominated
by the number of individuals in the Aufwuchs sample, whereas bdelloids were
dominant in the plankton sample.
The 7 monogononts found in the Aufwuchs sample were represented by the
Lepadellidae, Dicranophoridae, Lindiidae and Proalidae, with Lindia elsae sp. nov. dominant and Colurella
adriatica codominant. The
plankton sample showed 4 monogonont species only, belonging to the
Notommatidae, Lepadellidae and Synchaetidae, with an absolute dominance of Synchaeta squamadigitata sp. nov.
Seven of the monogonont species found are hitherto undescribed,
but 2 of them only, 1 species each of Lindia
and Synchaeta, occurred in sufficient
numbers to warrant description. The
description of the new species and comments on the other taxa are given here.
Table 1. Rotifera found in Aufwuchs (1) and plankton (2) samples
|
Sample |
1 |
2 |
|
Bdelloidea indet. |
|
31 |
|
Cephalodella
sp. |
- |
1 |
|
Colurella adriatica Ehrenberg, 1831 |
13 |
- |
|
C. colurus
Ehrenberg, 1830 |
- |
1 |
|
Encentrum marinum (Dujardin, 1841) |
1 |
- |
|
Encentrum
sp. 1 |
1 |
- |
|
Encentrum
sp. 2 |
2 |
- |
|
Lindia elsae
sp. nov. |
22 |
- |
|
Proales similis de Beauchamp, 1907 |
1 |
- |
|
Proales
sp. |
8 |
- |
|
Synchaeta squamadigitata sp. nov. |
- |
20 |
|
Synchaeta
sp. |
- |
2 |
a Number of specimens.
Lindia elsae sp. nov.
Differential diagnosis: Lindia elsae sp. nov. is
diagnosed by the epipharynx consisting of a single element with a large central
opening, showing 2 lateral extensions and 2 serrate semicircular parts. As such, it is easily separated from all
other known Lindia-species.
Material examined: Holotype a parthenogenetic female on a permanent, glycerin glass slide
mount deposited in the Koninklijk Belgisch Instituut voor Natuurwetenschappen
(KBIN),
Type locality: Infralittoral
fringe of Grande Anse, Grand Bois,
Etymology: The
species is dedicated to Dr. Els Van Rompu, in recognition of a long and
much-appreciated collaboration.
Description:
Parthenogenetic female (Figs. 2-5). Body of slightly
contracted specimens elongate, spindle-shaped (Figs. 2, 3). Head large, with 2 or 3 transversal
folds dorsally, rostrum broadly rounded.
Dorsal antenna in distal 1/3.
Auricles short-stalked.
Figs. 2-8. Lindia elsae sp. nov. (2) Lateral view; (3) dorsal view; (4)
toe, dorsal view; (5) toes, lateral view; (6) subitaneous egg; (7) epipharynx,
frontal view; (8) trophi, ventral view.
Scale bars: 2, 3, 6: 50 µm; 4, 5, 7, 8: 10 µm.
Trophi (Figs 7-15). Rami lyrate with distinct alulae;
basifenestrae large; inner margin of ramitips with scleropili (Fig. 11; rs); no
distinct external lamellae. Fulcrum
fairly long, ~1.5× ramus length, in lateral view almost parallel-sided and
slightly rounded distally. Unci with
4 (right) and 5 (left) rod-shaped and webbed teeth, gradually decreasing in
size, ventral teeth longest; dorsal teeth joined distally to a large triangular
web connected to the manubrium (Fig. 12; w); subunci a mass of short and small
scleropili (Fig. 11; su); outer side of ventral margin of major teeth with
preuncinal teeth, composed of a small irregular sclerite element and a larger,
rod-shaped, moveable element (Figs 11, 12, 15; pu). Ventral branch of manubria slender,
short, less than 1/2 length of shaft and nearly parallel to it; inner side of
head with small opening at base of the major tooth (Fig. 15; mo), a larger
ventral opening at turn between shaft and ventral branch (Fig. 9; vo), and a
large, funnel-shaped dorsal opening leading to a thin rib, running along greatest part of shaft
(Fig. 9; do). Epipharynx (Figs 7,
10, 13, 14) a single piece consisting of a central more or less flattened
O-shaped part, bearing 2 long lateral extensions and 2 serrate (~14 teeth)
semicircular dorsal extensions terminating in a smooth-margined and free
projection. It is shown in a
latero-lateral plane in the figures, but is actually oriented dorsoventrally in
the body, the lateral extensions pointing laterocaudally and the serrate
elements pointing rostrally. Semicircular
elements showing asymmetry: right one inserted near base of O-shaped part, and
left one inserted at base of lateral extension; both elements curving to right
side. Apparently, the epipharynx
situated around buccal tube.
Figs. 9-15. Lindia elsae sp. nov., SEM photographs
of trophi. (9) Dorsal view; (10)
ventral view; (11) detail of rami and unci tips; (12) unci, outer view; (13)
epipharynx, detail; (14) epipharynx; (15) uncus, inner view. do, dorsal opening; mo, opening at base
of major tooth; pu, preuncinal tooth; rs, rami scleropili; su, subuncus; vo,
ventral opening; w, web. Scale
bars: 9, 10, 12-15: 10 µm; 11: 5 µm.
Measurements: Total
length (n = 10) 175~205 (mean, 212) µm, toe 13~21 (mean, 16) µm; trophi (n = 8): ramus 8.7~12
(mean, 11.0) µm, fulcrum 13.3~16.4 (mean, 15.2) µm, uncus 8.0~9.6
(mean, 8.8) µm, manubrium 22.4~25.2 (mean, 24.1) µm, epipharynx (h × w) 32.4~37.4
× 8.4~12.7 (mean, 34.5 × 10.2) µm.
Comments:
The Lindiidae is a small family actually comprising 14 valid morphospecies
according to a recent revision by Segers (2002). It is characterized by the unique
cardate trophi which are modified for pumping. The body is illoricate with weakly
differentiated external morphology, and as such the species are very
similar. External characters of Lindia elsae sp. nov. that may aid in
differentiating it from some of the other Lindia
spp. are the single foot pseudosegment and the shape of the toes. Descriptors of the internal organization
allowing for some discrimination are the hyaline retrocerebral sac with few
red-pigmented granules, the club-shaped pedal glands with reservoirs, and
particularly the wide, transversally striated esophagus. Apparently, an annulated esophagus is
only shared with L. torulosa Dujardin,
1841. However, all species can be
recognized unequivocally by their complicated trophi, in particular the
characteristic epipharynx. Among
the Lindia spp. known to date (see
Segers 2002), the new species shows the relatively longest fulcrum. Lindia elsae sp. nov. and L. tecusa Harring and Myers, 1922 are
the only species displaying a straight anterior manubrial branch which is
almost parallel to the shaft, while the others have a crescent-shaped or
irregular (L. candida Harring and
Myers, 1922) branch. The new
species, however, is easily distinguished from L. tecusa by its different epipharynx (a broad plate without a
central hole and without serrate elements in L. tecusa), the conical
toes (toes very short, broadly conical and slightly compressed laterally in L. tecusa), the normal auricles (absent
from L. tecusa), and its
ovoviviparity (viviparity in L. tecusa). Serrate epipharynx elements have also
been demonstrated in L. torulosa (see
De Smet
Distribution and ecology: The new species is known from the type locality only. It was the dominant rotifer in Aufwuchs
among a stand of the brown alga Padina pavonia from the infralittoral
fringe. Examination of the gut
contents revealed that L.
elsae sp. nov. feeds on filamentous cyanobacteria, such
as Oscillatoria
sp. The new species was accompanied
by a rich fauna, consisting of foraminiferans, nematodes, polychaete annelids,
arthrotardigrades, kinorhynchs, copepods, ostracods, amphipods, molluscan
larvae, and other rotifers (Table 1).
To date, only 2 Lindia species have been reported from thalassic environments: L. tecusa
is an inhabitant of marine and brackish waters (Segers 2002), and L. torulosa,
although primarily a freshwater species, has been mentioned occasionally from
thalassic waters as well (Remane 1929, Schwarz 1955/56 1962, Saunders-Davies
1995 1998, De Smet 2002a ).
Synchaeta squamadigitata sp. nov.
Differential diagnosis: Synchaeta squamadigitata sp. nov. is diagnosed by its distally broadly rounded and squamate toes,
with proximally displaced ventrolateral tubuli. It cannot be mistaken for any of the
other known species of the genus.
Material examined: Holotype a parthenogenetic
female on a permanent, glycerin glass slide mount deposited in the KBIN,
Type locality. Port Est, Réunion I., Indian Ocean. 4 Jan. 2002.
Etymology: The
specific name squamadigitata (Latin squama meaning scale, digitata
meaning provided with toes) refers to the characteristic shape of the toes.
Description: Parthenogenetic female (Figs 16-19). Body more or less vasiform, neck region
narrower with 2 small transversal folds distally, and 2 dorsolateral humps
proximally; head almost 1/2 total length; trunk with a few weak dorsal
longitudinal striae. Lateral antennae
in posterior 1/3 of trunk. Apical
field strongly convex, with 2 small frontal antennae, and 2 small dorsal and
lateral coronal antennae bearing small styli. Auricles medium large, triangular, with
pronounced lateral tips recurved apically.
Dorsal antenna in neck region, on shallow prominence. Two fused, red cerebral eyes of equal
size (Fig. 19); apparently no frontal eyespots or granules. Foot medium long, conical, 4
pseudosegments. Toes double (Figs
17, 18), of equal length, short, basal part conical, distal part squamate with
broadly rounded margin; tubuli ventrolateral and subterminal. Pedal glands double, as long as foot,
with reservoirs in the toes.
Figs. 16-20. Synchaeta squamadigitata sp. nov. (16) Dorsal view; (17) foot, ventral
view; (18) toes, lateral view; (19) cerebral eyes; (20) trophi. Scale bars: 16: 50 µm; 17-20: 10 µm.
Trophi (Figs
20-28) weakly sclerotized. Unci
composed of a rather small frontal hook, and a toothplate bearing 5~6/6~7 blunt
teeth gradually decreasing in size; sulci between teeth shallow; gap between
frontal hook and toothplate fairly small, with membraneous web composed of tiny
scleropili; basal spine relatively small, basally with a small triangular and
inwardly projecting platelet composed of ~6 scleropili (Fig. 26; ss). Rami lamellar, composed of an elongate,
more-heavily sclerotized triangular inner lamella, and a weakly sclerotized
outer lamellar part; outer part composed of narrow median part, and large, rounded
outer part separated by a weakly reinforced ridge connected to toothplate;
proximal margin of rami with reinforced ridge; distal margin of elongate
triangular plate concave; inner margin of rami distally with row of scleropili
increasing in length distally.
Epipharynx (Fig. 25) a single element, consisting of 2 elongate semilunate parts, pointed
acutely on both ends, and a semicircular band connected to the semilunate parts
by a thin web. Fulcrum long, plate-shaped,
in lateral view slightly widening distally; distal margin oblique. Manubria long, shaft very weakly curved,
almost straight, anterior 1/3 with large, blunt triangular dorsal lamella;
anterior 1/2 with large semilunate ventral lamella; at anterodorsal margin of
dorsal lamella a series of scleropili forming a small triangular lamella (Fig.
21).
Figs. 21-28. Synchaeta squamadigitata sp. nov., SEM
photographs of trophi. (21) Right
manubrium, inner view; (22) incus, ventral view; (23) left manubrium, outer
view; (24) incus, dorsal view; (25) epipharynx; (26) detail of basal spine and
platelet; (27) unci, epipharynx, detail; (28) frontal view. Scale bars: 21-25, 27, 28: 10 µm; 26: 1
µm. bs, basal spine; fh, frontal
hook; rs, rami scleropili; ss, platelet at basal spine.
Measurements: Total length (n = 10) 98~138 (mean, 118) µm, toe 6~8 (mean, 7) µm; (n = 5) trophi 38~43 µm: ramus ~14~17 µm,
fulcrum 19~22 µm, uncus ~8~9 µm × 12~13 µm, manubrium 24~29 µm, epipharynx (h ×
w) 9~10 µm × 14~18 µm.
Comments: Synchaeta
squamadigitata sp. nov. is one of the smallest
members of the genus. Applying the
key in Hollowday (2002), the new species keys out near S. glacialis
Smirnov, 1932 and S.
hyperborea Smirnov, 1932, both of which are known from
Arctic seas, with which it only shares small coronal styles and a large
head. The other external features
of S. squamadigitata greatly differ, and its trophi also show little resemblance (Friedrich
and De Smet 2000). It is difficult
to relate S.
squamadigitata to any other Synchaeta on
the basis of the trophi morphology, since most of the available data are
unreliable and incomplete.
Of the genus Synchaeta, family Synchaetidae, some 34
species are recognized as being valid (Hollowday 2002). Diagnosis and identification
traditionally rely on the shape of the body and apical field; the position of
the lateral antennae; the shape of the foot and the number and shape of the
toes; the presence, type and number of eyespots; and to a lesser extent on the
trophi, especially the unci. Many
of these external distinguishing features appear variable or at least difficult
to ascertain. The trophi are
delicate and often fairly hard to see with LM, and little attention was paid to
their detailed structures in the past.
However, it is my experience that LM in combination with SEM can reveal
major taxonomically useful characters.
Of special interest are the unci, epipharynx, manubria (shaft straight
or kinked; the shape of the inner and outer lamella, their extent and the
position of their greatest width in relation to the shaft; and the shape of the
accessory lamella), and the rami triangular lamellae (concave or convex
anteriorly).
Identification
of this illoricate genus is considered virtually impossible in the preserved
state (e.g. Ruttner-Kolisko 1974, Rougier et al. 2000). However, they easily can be prepared and
preserved fully extended, using cocaine or eucaine (Rousselet 1902), procaine
hydrochloride (May 1988), or some local anesthetic (pers. observ.). Nogrady and Rowe (1993) obtained very
well anesthetized specimens with the addition of 20% (v/v) carbonated water to
the samples.
Distribution and ecology: Known from the
type locality only. It occurred in the plankton of the littoral zone
consisting, among others, of tintinnids; dinoflagellates; copepods; podonid
cladocerans; larvaceans; polychaete, molluscan, and echinoderm larvae; and
other rotifers (Table 1).
Comments on Other Species
Cephalodella sp.
A single
specimen (Figs 29, 30, 41) of an unknown Cephalodella
was found in the plankton sample.
The species is characterized by the finely denticulate (~22 spinules)
ventral margin of the slender, ventrally curved medium long toes. The trophi (Fig. 41) show a long fulcrum
and slightly asymmetrical manubria (with the left one larger) with axe-shaped
cauda.
The only other Cephalodella bearing spinulets at the
ventral margin of the toes, is the Tasmanian endemic C. lindamaya Koste and Shiel, 1986 described from acidic
freshwater. It is easily
distinguished from the species of Réunion by the simply incurved manubrial
cauda.
Figs. 29-33.
Habitus of rotifer species. (29) Cephalodella
sp.; (30) same as 29, detail toes; (31) Colurella
adriatica; (32) Proales sp.; (33)
Synchaeta sp. Scale bars: 10 µm.
Colurella spp.
A single
specimen of C. colurus was found in
the plankton; C. adriatica was relatively
numerous in the Aufwuchs. Both
species are euryhaline and eurytopic, and considered cosmopolitan. Several records exist from the Southern
Hemisphere (e.g. de Beauchamp 1913, Russell 1956, De Ridder and Segers 1997),
but it was not previously reported from marine habitats.
In C. adriatica different forms have been
described, with increased lorica size and toe length correlated with increasing
salinity (Hauer 1925), or with animals inhabiting running water showing a wider
lorica than those from stagnant biotopes (Donner 1964). However, these presumed relationships
between external morphology and environmental parameters remain to be proven by
thorough morphometric analyses. The
specimens encountered in the marine littoral of
Figs. 34-37. Colurella adriatica, SEM photographs of
trophi. (34) Frontal view; (35)
incus, frontal view; (36) caudal view; (37) lateral view. Scale bar: 10 µm.
Encentrum spp.
The genus Encentrum was represented by E. marinum and 2 undescribed species. Encentrum marinum (trophi in Fig. 38) is
a widespread species (De Ridder and Segers 1997, De Smet 1997), mainly known
from thalassic and athalassic waters, but also recorded from alkaline fresh
waters. Southern Hemisphere records
exist from saline ponds in
The undescribed
Encentrum sp. 1 (trophi shown in Fig.
39) is characterized by trophi bearing a shallow, broad tooth at the inner
margin of both rami, and a short, truncate-triangular fulcrum.
Encentrum sp. 2 (trophi shown in Fig.
40) has an elongate hexagonal rami outline, with the right ramus showing a
pronounced, distally pointing ventral extension. The species was also found in
interstitial sand of Peter the Great Bay and Ussuri Bay, Sea of Japan, Russia
(De Smet and Chernyshev unpubl. data), and thus is probably widespread.
Proales spp.
A single
specimen of P. similis (trophi shown
in Fig. 44) was found in Aufwuchs.
The species is known from thalassic and athalassic waters from most
parts of the world (De Smet 1996, De Ridder and Segers 1997). In the Southern Hemisphere, it is
recorded from inland saline waters in W. Australia (Bērziņš 1953),
The Aufwuchs
sample contained another and yet undescribed Proales sp. (Fig. 32), characterized by the toes with long,
needle-shaped points, reminiscent of the much larger P. syltensis Tzschaschel, 1978. It is distinctly differentiated from the
latter by the trophi (Figs 42, 43): the smaller manubrial lamellae, the large
basal apophyses, and the absence of large frontal rami plates.
Synchaeta sp.
Two specimens (Fig. 33) of
this undescribed species were found in the plankton sample. The apical field is strongly convex,
with 2 weak coronal styles; the body is cylindrical and slender; the foot is
short, with 2 equal toes; there is a single eyespot, located cerebrally. Trophi (Figs 45-48) strongly
sclerotized. The uncus plate has 2 or 3 fairly strong teeth and 3 shallow
smaller teeth, without a gap; the lamella in front of the teeth is strongly
sclerotized; the triangular lamella is concave distally; for the manubria, the
shaft is slightly curved, the anterior 1/2 has a large semilunate ventral
lamella, and rounded-triangular dorsal lamella; the greatest widths of the
lamellae are at ~1/4 and 1/7 from the proximal end respectively; an accessory
lamella is apparently absent; the epipharynx is composed of 2 elongate
semilunate elements connected by a semicircular element, and the semilunate
elements have openings proximally.
Figs. 38-48.
SEM photographs of trophi.
(38) Encentrum marinum, dorsal
view; (39) Encentrum sp. 1, ventral
view; (40) Encentrum sp. 2, ventral
view; (41) Cephalodella sp., lateral
view; (42,43) Proales sp. (42)
frontal view; (43) same as 42, detail of incus; (44) Proales similis, ventral view; (45-48) Synchaeta
sp. (45) epipharynx; (46) incus; (47) unci, frontal view; (48) right manubrium,
outer view. Scale bars: 10 µm.
Concluding Remarks
Twelve rotifer taxa (1
Bdelloidea undetermined and 11 Monogononta) are registered from Aufwuchs and
plankton of the marine littoral of tropical Réunion I. Both species richness and abundance were
low, which agrees with the very few observations from the different climatic
belts of the Northern and Southern Hemispheres. Although these are results of a very
limited sampling effort, the find of
7 undescribed species, two of which are described in the present paper,
and the 1st record of 2 cosmopolitan eurytopic species (Colurella adriatica and C.
colurus) from marine habitats in the Southern Hemisphere, once more
demonstrates the knowledge gap in marine rotifers. The presence of Bdelloidea likewise
shows that, although this group is considered primarily limnic and
limno-terrestrial, a yet-unexplored marine bdelloid fauna does exist. It is also clear that, even though the
marine rotifer fauna may be considered impoverished relative to its freshwater
counterpart, marine rotifer diversity is obviously underestimated, and will
ultimately turn out to be much richer considering the high diversity of marine
ecosystems. This lack of knowledge
must be ascribed to difficulties in obtaining well-preserved specimens, the low
densities at which most of the species occur, the lack of taxonomic expertise,
and identifications to higher taxonomic levels only by ecologists, and the few
taxonomists working on rotifers, especially marine ones.
Acknowledgments: The samples
were taken at the occasion of expeditions to the Subantarctic that were
partially funded by the Science Foundation Flanders (FWO). Dr. Bart Vandevijver and Drs. Pieter Ledeganck are acknowledged for collecting the
Aufwuchs sample. The Laboratory of
Cell Biology and Histology, University
of Antwerp
----------------------------------------------------------------------------------------------------------------
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