J OURNAL OF C RUSTACEAN B IOLOGY, 32(5), 871-882, 2012

A NEW PHREATIC SPECIES OF THE GENUS PARVULOBATHYNELLA

(MALACOSTRACA: BATHYNELLACEA) FROM SOUTHEASTERN INDIA, ALONG
WITH AN UPDATED KEY TO THE SPECIES

Yenumula Ranga Reddy ∗ and Venkateswara Rao Totakura ∗∗

Department of Zoology, Acharya Nagarjuna University, Nagarjunanagar 522 510, India

ABSTRACT
Up until now, the genus Parvulobathynella Schminke, 1973, had eight species in the world. Of these, three species each are known from
South America and Africa, and two from peninsular India. Parvulobathynella macrodentata n. sp. is described herein from the phreatic
zone of southeastern India. Interestingly, the new species has striking mandibular affinity with the Brazilian Odontobathynella amazonica
Delamare Deboutteville and Serban, 1979. So the mandibular criteria of Parvulobathynella are suitably amended to accommodate the new
species and the interrelationships of Parvulobathynella and Odontobathynella, briefly discussed. The paper also provides an updated key
to the world species of Parvulobathynella in addition to some ecological notes on the new species.
K EY W ORDS: Bathynellacea, India, Parvulobathynella, stygofauna, taxonomy
DOI: 10.1163/193724012X645844

I NTRODUCTION M ATERIAL AND M ETHODS

We collected the specimens we studied by filtering the
Parvulobathynella Schminke, 1973, has recently been re- groundwater when it was being pumped out of farm bores
vised by Ranga Reddy et al. (2011). Today, it is rep- (depth c. 10 m). Filtering was done manually by holding
resented by eight species in the world, which are all a bolting-silk plankton net (mesh size 70 μm) against the
distributed on Gondwanan fragments: Parvulobathynella water current for 20-30 minutes at each time of sampling as
ypacaraiensis (Noodt, 1963), P. camposicola (Jakobi, 1961), pictured in Ranga Reddy and Totakura (2012). The filtrate
and P. riegelorum (Noodt, 1965) in South America, P. was fixed in 5% formaldehyde.
Specimens were sorted into 70% alcohol and later trans-
pentodonta (Serban and Coineau, 1982), P. duodecima Cho
ferred into glycerol. Dissection was carried out in glycerol
and Schminke, 2001, and P. octacantha Cho and Schminke, under a binocular stereomicroscope at a magnification of
2001 in Africa, and P. distincta Ranga Reddy et al., 2011, 90×. Drawings were made with the aid of a drawing tube
and P. projectura Ranga Reddy et al., 2011 in southeastern mounted on a Leica DM 2500 Trinocular Research Micro-
India. Morphologically, the species constitute an extremely scope, equipped with UCA condenser, IC objective prism
homogeneous group and most of them live in hyporheic and 1-2× magnification changer. Permanent preparations
habitats. were mounted in glycerol. The type material was deposited
During our on-going investigations on the Indian sty- in the Muséum national d’Histoire naturelle, Paris.
gofauna, we have encountered a new species, Parvulo-
bathynella macrodentata, in the phreatic waters of south- S YSTEMATICS
eastern India. This is a remarkable species in that its Order Bathynellacea Chappuis, 1915
mandibular details alone are almost typical of the Brazil- Leptobathynellidae Noodt, 1965
ian Odontobathynella amazonica Delamare Deboutteville Parvulobathynella Schminke, 1973
and Serban, 1979. Accordingly, the mandibular criteria of
Amended Diagnosis.—See Discussion below.
Parvulobathynella are further amended to accommodate this
species. The paper gives an illustrated description of the Parvulobathynella macrodentata n. sp.
new species together with a brief discussion on the inter- Type Locality.—Farm bore, Peravaram village
relationships of Parvulobathynella and Odontobathynella. (16°53 42.0 N, 81°45 09.8 E; elevation 17.7 m; water tem-
Furthermore, Schminke’s (2011) key to species of Parvu- perature 26°C; pH 7.0), ∼20 km from Rajahmundry, East
lobathynella is updated by including the Indian congeners. Godavari District, Andhra Pradesh, South India, Coll. V. R.
A few ecological notes on the new species are also added. Totakura, 12 April 2008.

* Corresponding author; e-mail: [email protected]

** E-mail: [email protected]

© The Crustacean Society, 2012. Published by Brill NV, Leiden DOI:10.1163/193724012X645844

872 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 32, NO. 5, 2012

Fig. 1. Map showing the distribution of Parvulobathynella macrodentata n. sp. ( type locality;  other localities).

Material Examined.—Holotype male, dissected on 6 slides lateral arms. Labrum with 6 backwardly directed spinules
(MNHN-IU-2009-1703); allotype female, dissected on 5 and 4 dentate structures on a linguiform lobe on each side.
slides (MNHN-IU-2009-1704); 1 male paratype, whole- Distal endite of maxillule stumpy. Th I and II without
mounted on a slide (MNHN-IU-2009-1705); deposited in epipod. Uropodal sympod 3.3 times as long as wide, with
the National Museum of Natural History, Paris. 2 relatively large, equal spines; endopod 2.7 times as
Other Localities: Farm bore at Ravulapalem village long as wide, with 5 armature elements including 1 large
(16°06 33.4 N, 81°46 49.9 E; elevation 37 m; depth c. subterminal inner spine. Female Th VIII greatly reduced;
10 m; water temperature 27°C; pH 7.0), ∼20 km from Ra- male Th VIII with 1 seta postero-distally.
jahmundry, East Godavari District, Andhra Pradesh, South
India, Coll. V. R. Totakura, 12 April 2008. One male para- Description of Adult Male (Holotype).—Total length
type (dissected on 7 slides). Farm bore at Mamillapalli vil- 0.67 mm. Body slender, elongate, 12.5 times longer than
lage near Nidubrolu (16°02 23.8 N, 80°32 35.4 E; elevation maximum width; cuticle thin and imperforate. In lateral view
32 m), Guntur District, Andhra Pradesh, South India, Coll. (Fig. 2A), pleomeres about as wide as thoracic segments.
K. Yesubabu, 01 February 2011. One male paratype (dis- Head 1.3 times as long as wide and equaling the combined
sected on 3 slides); all in V. R. Totakura’s collection. length of first 2 thoracic segments.
Antennule (Fig. 3A): 6-segmented, 16% longer than head.
Diagnosis.—Antennular segment 5 with 1 large aesthetasc Length of first 3 segments 2.1 times as long as last 3
and segment 6 with 3 unequal subterminal aesthetascs. segments. Segment 1 with 2 plumose setae on outer distal
Molar teeth large, claw-like and contiguous at base; ventral- margin and 1 plumose and 2 simple setae on dorso-distal
edge tooth about as large as molar teeth and with a denticle surface. Segment 2 with 1 ventro-distal plumose seta close
at base; pars incisiva produced and larger than ventral- to outer margin, 3 plumose setae on distal margin in a
edge tooth, proximal-most tooth horizontal to other teeth. row, and 1 simple seta at inner distal corner. Segment 3
Paragnaths with 2 large papillate structures and massive with 1 long simple and 1 plumose setae at outer distal
 RANGA REDDY AND TOTAKURA: NEW PHREATIC SPECIES OF THE GENUS PARVULOBATHYNELLA 873

Fig. 2. Parvulobathynella macrodentata n. sp. A-B, paratype male. A, habitus, lateral view; B, head, ventral view.

corner. Inner flagellum short, rectangular, with 3 unequal curved and backwardly directed spinules and 4 dentate struc-
simple setae. Segment 4 with stout apophysis overreaching tures on large linguiform lobe on each side; medial spin-
midlength of next segment and bearing 2 slightly subequal ules longer than lateral ones; 2 small teats on ventral side
plumose setae. Segment 5 with 1 simple seta and 1 large (Fig. 3C).
aesthetasc at outer distal corner, 1 long simple seta on dorso- Mandible (Figs. 2B, 4A-D): pars incisiva with 4 teeth,
distal margin, 1 simple and 1 plumose setae at inner distal proximal-most tooth smallest, parallel to other teeth and
corner. Segment 6 with 4 unequal simple setae and 3 unequal close to palp (in situ, incisor process of right mandible
subterminal aesthetascs. perfectly fitting into the space between incisor process and
Antenna (Figs. 2B, 3B): 5-segmented, bent backwards
ventral-edge tooth of left mandible as in Fig. 2B). Tooth
between segments 1 and 2, extending to about midlength
of ventral edge strongly developed, somewhat shorter than
of head, in lateral view (Fig. 2A) and reaching 72.1%
of antennular length. Combined length of segments 1-3 pars incisiva and with 1 proximal denticle. Pars molaris
equaling that of segments 4-5. Segment 1 bare. Segment 2 consisting of 3 large, claw-like teeth, lying contiguous to
armed with 1 outer seta. Segments 3 and 4 subequal in each other at base and not far from tooth of ventral edge;
length, each with 1 long simple seta near inner distal molar teeth fringed with long spinules on lateral margins,
corner. Segment 5 with 2 terminal simple setae and 1 thick but spinules discernible only in certain angles of view. Palp
subterminal plumose seta. inserted at the level of ventral-edge tooth, 1-segmented,
Labrum (Figs. 2B, 3C): small, somewhat spherical in out- slightly swollen at base, almost twice as long as wide and
line; free margin slightly wavy, carrying 6 long, somewhat with 1 long simple seta.
874 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 32, NO. 5, 2012

Fig. 3. Parvulobathynella macrodentata n. sp. A-C, holotype male; D, allotype female. A, antennule, dorsal view; B, antenna, dorsal view; C, labrum,
ventral view; D, same, ventro-frontal view (in situ).

Paragnaths (Figs. 2B, 5A): represented by 2 large papillate spiniform seta arising from small protuberance on inner
structures; lateral arms massive. proximal margin. Third segment very small, rectangular,
Maxillule (Fig. 5B): proximal endite small, slightly longer with prehensile apical claw; claw 1.2 times longer than
than wide, with 4 similar claw-like spines at inner distal cor- second segment, with slightly swollen proximal third and
ner. Distal endite stumpy, somewhat rectangular in outline, very finely serrulate distal inner margin.
1.5 times as long as wide, with 3 apical and 2 inner claws Thoracopods I-VII (Figs. 6A-D, 7A-C): well developed,
and 2 simple setae on outer distal margin. length gradually increasing from Th I-IV; Th V-VII almost
Maxilla (Fig. 5C): 3-segmented. First segment nearly similar in size. Th I-II without epipod. Th III-VII each bear-
rectangular and unarmed. Second segment 2.6 times as ing 1-segmented, club-shaped epipod, 0.6 times as long as
long as wide and armed with 9 setae, of which 1 short, basis. Basis without inner marginal seta on Th I-VII. Exopod
 RANGA REDDY AND TOTAKURA: NEW PHREATIC SPECIES OF THE GENUS PARVULOBATHYNELLA 875

Fig. 4. Parvulobathynella macrodentata n. sp. A-C, holotype male; D, allotype female. A, mandible, ventral view; B, same, dorso-ventral view; C, same,
frontal view; D, same, nearly frontal view.

2-segmented, as long as endopod on Th I and II and 1.2 times 1 subterminal plumose setae and 1 row of spinules at base of
as long as endopod on Th III-VII. On all thoracopods, seg- subterminal plumose seta. Endopod 4-segmented; segments
ment 1 of exopod with 1 barbed seta on inner distal margin 2 and 3 with 1 row of spinules each on distal inner margin;
and 1 row of spinules; segment 2 with 1 terminal barbed and setal formula: 0 + 0 / 0 + 1 / 0 + 0 / 2(1).
876 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 32, NO. 5, 2012

Fig. 5. Parvulobathynella macrodentata n. sp. B, C, holotype male; A, D, allotype female. A, paragnaths, ventral view; B, maxillule, lateral view; C,
maxilla, lateral view; D, Th VIII, ventral view.

Thoracopod VIII (Fig. 8A-E): small, somewhat obovate, Uropod (Fig. 9B): sympod 3.3 times as long as wide
1.3 times as long as wide. Protopod massive. Outer lobe in dorsal view, bearing 2 similar, relatively long serrulate
small, conical, fused with penile lobe, but not discernible spines on slightly dilated inner distal margin. Exopod almost
in lateral and ventral views. Inner lobe shorter than dentate cylindrical, 3.3 times as long as wide, measuring 60%
lobe. Dentate lobe only with 2 denticles. Basis not defined of sympod length and armed with 1 long terminal and 1
at base and with 1 subdistal seta at base of exopod, perhaps short subterminal barbed setae. Endopod nearly 2.7 times
representing endopod. Exopod distinct, hook-like, apically as long as wide, 63% of sympod length, somewhat dilated
denticulate plate. subproximally and fringed with 2 crescentic rows of spinules
Pleopod 1: absent. on inner margin; armature consisting of 1 dorso-medial
 RANGA REDDY AND TOTAKURA: NEW PHREATIC SPECIES OF THE GENUS PARVULOBATHYNELLA 877

Fig. 6. Parvulobathynella macrodentata n. sp. A-D, holotype male, Th I-IV.

plumose seta, 2 barbed setae (1 terminal and 1 outer 2 dorsal plumose setae, of which outer seta 4.1 times as
subterminal; terminal seta 0.4 times as long as subterminal long as inner one; 1 row of fine spinules on ventro-distal
seta), 1 greatly reduced terminal seta and 1 large, proximally margin.
dilated, bipinnate spine at subterminal inner angle.
Description of Adult Female (Allotype).—Total length
Pleotelson (Fig. 9A): without any armature. 0.72 mm. Body and all appendages except labrum and tho-
Anal operculum (Fig. 9A): concave. racopod VIII as in male.
Caudal furca (Fig. 9A): somewhat rectangular, 1.6 times Labrum: frontal view (Fig. 3D) showing 4 incurved
as long as maximum width and rounded terminally, armed medial spinules in 2 pairs and 2 isolated lateral spinules, all
with 1 stout, proximally dilated, bipinnate terminal spine arising from small lobes, and 4 dentate structures on large
and 2 unequal spines (1 inner, subterminal, 1 dorsal) and dentate linguiform lobe on either side.
878 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 32, NO. 5, 2012

Fig. 7. Parvulobathynella macrodentata n. sp. A-C, holotype male, Th V-VII.

Thoracopod VIII (Fig. 5D): remarkably reduced, being D ISCUSSION

represented by thin crescentic lobe on either side of ventro- Generic Affiliation
median axis of the segment.
Judging by the mandibular morphology, which of course
provides vital supra-specific clues in bathynellacean taxon-
Variation.—Body size 0.61-0.67 mm in male. Number omy (Noodt, 1965; Schminke, 1972; Serban, 1980; Ranga
and size of spinules on labrum showing inter-population Reddy and Totakura, 2010), one might assign P. macroden-
variation (Fig. 3C, D). tata to the monotypic genus Odontobathynella rather than
to Parvulobathynella. The large, claw-like, contiguous mo-
lar teeth together with the likewise large, denticle-bearing
Etymology.—The specific epithet, macrodentata, alludes to ventral-edge tooth as well as pars incisiva of the new species
the unusually large mandibular teeth; derived from the Greek are almost identical to what obtains in the Brazilian O.
prefix macro = large, and the Latin adjective dentatus = amazonica. The heretofore known contrasting situation of
toothed. The name agrees in gender with the (feminine) Parvulobathynella consists primarily in the molar teeth be-
generic name. ing closely-spaced instead of contiguous and pars incisiva
 RANGA REDDY AND TOTAKURA: NEW PHREATIC SPECIES OF THE GENUS PARVULOBATHYNELLA 879

Fig. 8. Parvulobathynella macrodentata n. sp. Holotype male, Th VIII (abbreviations: dlb = dentate lobe; enp = endopod; exp = exopod; ilb = inner lobe;
olb = outer lobe). A, B, holotype; C-E, paratype. A, latero-external view; B, latero-internal view; C, anterior view; D, posterior; E, ventral view.

relatively small like the generally simple ventral-edge tooth. the true phylogenetic relationships of all these putatively dis-
In regard to the mandibular structure, Parvulobathynella is tinct genera.
rather closely related to Leptobathynella Noodt, 1965 and Regarding the principal non-mandibular generic criteria,
Califobathynella Cho, 1997. On the other hand, Odonto- however, Parvulobathynella is quite distinct from Odonto-
bathynella, as already emphasized by Delamare Debout- bathynella: 1) the free margin of labrum is generally spin-
teville and Serban (1979) and others, comes closest to the ulose vs. dentate; 2) the proximal endite of maxillule has
monotypic Braslibathynella Jakobi, 1958, in having contigu- four vs. two armature elements; 3) the male Th VIII has
ous molar teeth and large ventral-edge tooth, but differs from one seta vs. two claw-like setae in the postero-distal region;
the latter by having large vs. small pars incisiva and three vs. 4) the caudal furca has a simple vs. complex apical spine;
four molar teeth. Further research is necessary to understand and 5) the uropodal exopod has two setae (proximal seta ab-
880 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 32, NO. 5, 2012

Fig. 9. Parvulobathynella macrodentata n. sp. A-B, holotype male. A, pleotelson, dorsal view; B, uropod, dorsal view.

sent) vs. three setae (proximal seta present), and the endo- molar teeth; pars incisiva short or produced, proximal-most
pod has a total of five armature elements (inner subterminal tooth horizontal or perpendicular to other teeth.
spine present) vs. three armature elements (inner subtermi-
nal spine absent). Interspecies Relationships
Notwithstanding its apparent mandibular similarity with Clearly, P. macrodentata occupies the basal position in the
Odontobathynella, P. macrodentata is overwhelmingly close genus Parvulobathynella. A combination of the following
to the genus Parvulobathynella, fulfilling all other generic character states bear this out:
criteria of its recently amended diagnosis (see Ranga Reddy 1) Mandibular teeth are large relative to the corpus man-
et al., 2011). Thus, to accommodate the new species in the dibula; pars incisiva is particularly protruding whereas
it is short in other species except P. ypacaraiensis and
genus Parvulobathynella, we propose here a fresh amend-
P. riegelorum in which it is moderately developed.
ment to its generic diagnosis in regard to the mandibular cri- 2) Paragnaths are in the form of large papillate structures
teria alone as follows: molar teeth small spiniform or large and lateral arms massive. On the contrary, paragnaths
claw-like, contiguous or closely spaced; tooth of ventral are small protrusions and the lateral arms either
edge with or without proximal denticle and about as large as inconspicuous or much shorter than paragnaths in all
 RANGA REDDY AND TOTAKURA: NEW PHREATIC SPECIES OF THE GENUS PARVULOBATHYNELLA 881

other congeners. In P. ypacaraiensis alone, paragnaths 8. Uropodal endopod with small, naked, terminal spiniform
bear a “seta” each. seta; labrum with 8 spinules . . . . . . . . . . . . . P. octacantha
3) Distal endite of maxillule is stumpy instead of elongate – Uropodal endopod with tiny terminal spine; labrum with
or conical as in all other species. 28 spinules . . . . . . . . . . . . . . . . . . . . . . . . . . P. ypacaraiensis
4) The third maxillary segment is distinct as in the
African and South American congeners, whereas it Ecology
is fused with the claw in the two known Indian P. macrodentata is the second species of Parvulobathynella
congeners. A total of only nine setae occur on the distal reported from the phreatic waters, the first being the Brazil-
one or two segments as compared to ten setae in all ian species P. camposicola. All other species inhabit the
other congeners. sandy banks of rivers or, rarely, lakes, e.g., the Paraguayan
5) Labrum has a large linguiform lobe on either side, species P. ypacaraiensis. It is also noteworthy that all the
bearing four dentate structures. Whereas this lobe species except P. ypacaraiensis occur in the coastal belt.
(more clearly visible in frontal than ventral view) is P. macrodentata is rather rare in its occurrence. It was
inconspicuous or absent in the Indian congeners (P. found in farm bores, the depth of which is about 10 m. Water
distincta and P. projectura), it is a narrow bidentate temperature aboveground was 26-27°C and pH around 7.0.
structure in the others. It is observed that the number The new species occurs only with Serbanibathynella sp.
and size of the spinules are subject to inter-population, at Mamillapalli and with no other species at the other two
as already described (cf. Cho, 1995 in Cho, 1997). localities.
Another characteristic feature of Parvulobathynella is
that the spinules are always backwardly directed in
ventral view. Exceptionally, labrum is smooth in P. ACKNOWLEDGEMENTS
projectura. One of us (YRR) is grateful to the University Grants Commission, New
6) Th III has epipod. This character is shared only with P. Delhi, for providing financial support under the scheme of Emeritus Fellow-
ship. Material for this study was collected during 2008-2011 under a Ma-
riegelorum, P. duodecima and P. pentodonta jor Research Project (SR/SO/AS/25/2007) sanctioned by the Department of
Interestingly, the female Th VIII is greatly reduced in P. Science and Technology, Ministry of Science and Technology, Government
macrodentata as compared with its known congeners. On of India, New Delhi.
the other hand, it is untraceable in O. amazonica.
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