JOURNAL OF CRUSTACEAN BIOLOGY, 24(2): 246–260, 2004

A NEW GENUS AND SPECIES OF THE FAMILY DIOSACCIDAE

(COPEPODA: HARPACTICOIDA) FROM THE GROUNDWATERS OF INDIA

Tomislav Karanovic and Y. Ranga Reddy

(TK, correspondence) Western Australian Museum, Francis Street, Perth WA 6000, Australia
(karanovic@museum.wa.gov.au);
(YRR) Nagarjuna University, Department of Zoology, Nagarjunanagar 522510, Andhra Pradesh, India
(yrangareddy@yahoo.com)

ABSTRACT
The recent renewed interest in the Indian stygofauna has led to the discovery of another interesting
copepod representative, from the almost exclusively marine family Diosaccidae Sars, 1906. Neo-
miscegenus n. gen. and Neomiscegenus indicus n. sp., from the subterranean freshwaters of the Krishna
River at Vijayawada, Andhra Pradesh, are described. The new genus belongs to that evolutionary line of
Diosaccidae that encompasses the genera Amphiascoides Nicholls, 1941; Paramphiascella Lang, 1948;
Paramphiascoides Wells, 1967; and Miscegenus Wells, Hicks, and Coull, 1982. The genus Miscegenus
has the highest number of synapomorphies with Neomiscegenus, including the shape of the caudal rami
and the armature formula of the swimming legs. A peculiar shape of the genital field in the female and the
armature of the antennal exopod in both sexes are the most important autapomorphies of the new genus,
while the fine structure of the sexually dimorphic second leg in the male is probably plesiomorphic. The
phylogenetic position of the new genus is briefly discussed.

In a global context, the stygofauna of the Indian scribed as a new genus in the harpacticoid family
subcontinent is very poorly known (Marmonier Diosaccidae Sars, 1906.
et al., 1993; Ranga Reddy, 2002b, in press b), The family Diosaccidae presently encom-
with only a few records until the mid-1980’s passes about 388 species and subspecies,
(Botosaneanu, 1986). As far as the freshwater classified into 43 valid genera (Lang, 1948;
subterranean copepods are concerned (Lescher- Bodin, 1997; Mu and Gee, 2000). They are
Moutoué, 1986; Rouch, 1986), by that time only predominantly marine free-living forms, with
six species of the genus Parastenocaris Kessler, only a few species reported as commensals or
1913, from Sri Lanka (Enckell, 1970) and one semiparasites on lobsters (Humes, 1953). Two
species of the genus Elaphoidella Chappuis, large genera, Schizopera Sars, 1905, and
1928, from the mainland (Chappuis, 1954) had Stenhelia Boeck, 1864, represent almost 35%
been described. Four cyclopoids, reported by of the whole family, while even 22 genera
Pesce and Pace (1984) from several Indian (more than 50%) have three or fewer species,
freshwater wells, are actually all stygophiles (or and 12 genera (28%) are monospecific. Obvi-
even stygoxenes), not stygobites. The recent ously, this is not a young and aggressive family,
renewed interest in the Indian stygofauna has but rather just a remnant of the former, and
revealed a number of very interesting sub- probably very distant, flourishing. That may be
terranean copepods (Ranga Reddy, 2001; Kar- one of the reasons for their relatively unsuccess-
anovic and Pesce, 2001; Karanovic and Ranga ful colonization of the freshwater habitats,
Reddy, in press), as well as some ancient although the genus Schizopera does have
bathynellids (Ranga Reddy, 2002a; in press a). a number of freshwater forms, with a small
The latter probably emerged from their marine species flock of about 10 species in the ancient
ancestors as far back as the Permian (Schminke, Lake Tanganyika (Boxshall and Jaume, 2000).
1981; Pandourski and Ognjanova, 2001), i.e., But even those representatives found in inland
just after the Permo-Carboniferous glaciation, continental habitats prefer waters of increased
which occurred when India formed part of the salinity (Karanovic, 2004). However, because
Gondwana supercontinent (Frakes, 1999; Play- the stygal component of the freshwater copepod
ford, 2003). In this paper, another freshwater fauna is practically unknown in many parts of
stygobiont with clear marine ancestry is de- the world, and especially in the former parts of

246
 KARANOVIC AND RANGA REDDY: NEOMISCEGENUS, N. GEN., FROM INDIA 247

Gondwana, it is very hard to make even the trally in acuminate lappets; ancestral proximal
most general assumption about the freshwater lateral seta inserted ventrally at midlength of
colonizations. The new genus described here, as ramus; dorsal seta inserted medially, close to
well as another one described earlier (Karanovic inner margin. Female antennula about 1.5 times
and Pesce, 2001) from the almost exclusively as long as rostrum and clearly 8-segmented.
marine family Ectinosomatidae Sars, 1904, Antenna with allobasis and 3-segmented exo-
makes one just wonder what the Indian copepod pod; second exopodal segment unarmed, third
stygofauna looks like. One of the implications one armed with 1 apical and 1 lateral seta.
of this finding is the beginning of our realization Mandibula with 1-segmented endopod and exo-
that the colonization of the subterranean fresh- pod; exopod with 2 lateral and 2 apical setae.
water habitats may have a very different history Maxillula also with 1-segmented endopod and
in parts of the former Gondwana from that in the exopod; endopod armed with 1 lateral and 3
Northern Hemisphere. apical setae. Maxilla with 2, 2, and 3 setae on
syncoxal proximal, middle, and distal endites
MATERIALS AND METHODS respectively. Maxilliped with only 2 setae at
The sample was collected from the subterranean passages distal inner corner of basis. All swimming legs
flowing into the River Krishna near Vijayawada from the with 3-segmented exopods and endopods; first
adjacent, soaked, agricultural land. About a week before swimming leg with first endopodal segment half
sampling, an abnormal flood occurred in the river, inundat- as long as exopod; female swimming leg arma-
ing the low-lying areas of southern bank. The study area and
the recorded microfauna were described by Ranga Reddy ture formula same as in the genera Amphias-
(2001). A plankton net (mesh size 70 lm) was held against coides, Paramphiascella, Paramphiascoides,
the groundwater runoff for about 30 minutes. The material and Miscegenus. Female fifth leg with 5 strong
was preserved in 10% formaldehyde solution, and the cope- armature elements on baseoendopod and 5
pods were later separated with a dissecting microscope and
transfered to 70% ethanol. Specimens were dissected in
elements on exopod: middle one slender, seti-
Fauré’s medium, which was prepared following the proce- form, and apically inserted, others spiniform.
dure discussed by Stock and von Vaupel Klein (1996). Dis- Female sixth leg with 2 setae; outer one stouter
sected appendages were covered with a coverslip. For the and plumose. Male with sexually dimorphic
urosome or the whole animal, two human hairs were pore on third exopodal segment of third leg and
mounted between the slide and coverslip, so the parts could
not be compressed. By moving the coverslip carefully by with transformed second and third segments on
hand, the whole animal or a particular appendage could be second leg: ancestral outer apical seta on third
positioned in different aspects, making possible the observa- segment transformed into serpentine, smooth
tion of morphological details. During the examination, water and strong element, with swollen distal end and
slowly evaporated, and appendages or the whole urosome
eventually remained in completely dry Fauré’s medium. All
2 characteristic tips on it.
drawings were prepared using a drawing attachment (tube)
on a Leica DMLS brightfield compound microscope, with Type and Only Species.—Neomiscegenus indi-
C-PLAN achromatic objectives (403/0.65; 633/0.75 and
1003/1.25 oil). Morphological terminology follows Huys cus, n. sp.
and Boxshall (1991), except for the swimming legs armature
formula, where a simplified version is used. The material is Etymology.—The generic name is composed of
deposited in the Western Australian Museum, Perth (WAM). the genus name Miscegenus, which is morpho-
The new species was never found in the core samples col- logically closest to the new genus, and the Greek
lected during 1998–2001 in the riverbed. prefix ‘‘neo’’ (meaning: new). Gender masculine.

RESULTS Neomiscegenus indicus, n. sp.

Order: Harpacticoida Sars, 1903 Figs. 1–29
Family: Diosaccidae Sars, 1906
Neomiscegenus, n. gen. Material Examined.—Holotype: Female (WAM
C28608) - India, Andhra Pradesh, Krishna River
Diagnosis.—Small Diosaccidae with fusiform at Vijayawada, groundwater runoff on the
habitus. Female genital double-somite with rigid southern bank, 24 October 1998, leg. Y. Ranga
internal sclerotized ridge; genital complex with Reddy, 168319N 808409E: dissected on 2 slides.
single large copulatory pore and without epicop- Allotype: Male (WAM C28609) - India,
ulatory bulb, with short copulatory duct and 2 Andhra Pradesh, Krishna River at Vijayawada,
small, reniform, seminal receptacles. Caudal groundwater runoff on the southern bank, 24
rami cylindrical, more than 1.5 times as long as October 1998, leg. Y. Ranga Reddy, 168319N
wide, with posterior margins terminating ven- 808409E: dissected on 1 slide.
248 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 24, NO. 2, 2004

Figs. 1–6. Neomiscegenus indicus, n. gen., n. sp., holotype (female): 1 - habitus, dorsal view; 2 - antennula; 3 - left antenna;
4 - labrum; 5 - rostrum; 6 - exopod of right antenna. Scales ¼ 0.1 mm.
 KARANOVIC AND RANGA REDDY: NEOMISCEGENUS, N. GEN., FROM INDIA 249

Figs. 7–10. Neomiscegenus indicus, n. gen., n. sp., holotype (female): 7 - abdomen, ventral view; 8 - mandibula; 9 - coxa of
mandibula; 10 - anal somite and right caudal ramus, lateral view. Scale ¼ 0.1 mm.
250 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 24, NO. 2, 2004

Figs. 11–15. Neomiscegenus indicus, n. gen., n. sp., holotype (female): 11 - urosome, dorsal view; 12 - maxillula; 13 -
maxilla; 14 - maxilliped; 15 - fifth leg. Scale ¼ 0.1 mm.
 KARANOVIC AND RANGA REDDY: NEOMISCEGENUS, N. GEN., FROM INDIA 251

Figs. 16–17. Neomiscegenus indicus, n. gen., n. sp., holotype (female): 16 - first swimming leg; 17 - second swimming leg.
Scale ¼ 0.1 mm.
252 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 24, NO. 2, 2004

Figs. 18–19. Neomiscegenus indicus, n. gen., n. sp., holotype (female): 18 - third swimming leg; 19 - fourth swimming leg.
Scale ¼ 0.1 mm.

Description.—Female (holotype): Total body plius eye not visible. Habitus (Fig. 1) fusiform
length, measured from tip of rostrum to posterior but relatively slender, compressed dorsoven-
margin of caudal rami (excluding caudal setae), trally, without distinct demarcation between
0.417 mm. Preserved specimen yellowish. Nau- prosome and urosome; prosome/urosome ratio
 KARANOVIC AND RANGA REDDY: NEOMISCEGENUS, N. GEN., FROM INDIA 253

Figs. 20–29. Neomiscegenus indicus, n. gen., n. sp., allotype (male): 20 - pseudoperculum, dorsal view; 21 - abdomen,
ventral view; 22 - spermatophore; 23 - antennula; 24 - third exopodal segment of second swimming leg; 25 - third exopodal
segment of third swimming leg; 26 - endopod of second swimming leg; 27 - fifth leg; 28 - basis of first swimming leg; 29 -
sixth leg. Scales ¼ 0.1 mm.
254 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 24, NO. 2, 2004

1.2 and greatest width at second pedigerous (first width (without lappets); ornamented with many
free) somite. Body length/width ratio about 3.3; hairs along inner margins, 1 ventral cuticular
cephalothorax 1.36 times as wide as genital pore each, and 1 setula at base of lateral seta, as
double-somite. Free pedigerous somites without well as with posterior row of large spinules and
particular expansions laterally or dorsally. In- transverse row of several spinules ventrally,
tegument strongly chitinized, but with smooth between ventral seta and posterior outer corner;
surface. Rostrum (Figs. 1, 5) long, reaching armed with 6 setae, 1 ventrally, 1 laterally, 1
distal margin of third antennular segment, dorsally, and 3 apically. Dorsal seta about 1.5
linguiform, with blunt tip, about 1.7 times as times as long as ramus from dorsal view,
long as wide and clearly demarcated at base; inserted at midlength of ramus close to inner
ornamented with 2 sensilla laterally. Cephalo- margin, biarticulate at base and smooth. Ventral
thorax (without rostrum) slightly longer than seta (in fact ancestral lateral proximal seta,
wide, representing 33% of total body length. which moved strongly ventrally) smooth, in-
Surface of dorsal shield covering cephalothorax serted at middle of ramus and about 0.7 times as
with many large sensilla, as well as tergites of 3 long as ramus. Lateral seta (ancestral distal
free pedigerous somites. Hyaline fringes of all lateral one) also smooth and slightly shorter
prosomites narrow, smooth. Fifth pedigerous than ramus, inserted at 3/5 of ramus length.
somite (first urosomal) ornamented with 6 dorsal Inner apical seta bipinnate, 1.3 times as long as
sensilla and with hyaline fringe finely serrated. ramus, arising from small protuberance. Middle
Genital double-somite (Figs. 7, 11) about as long apical seta strongest, bipinnate at distal end,
as wide (ventral view), without subdivision line with breaking plane, but obviously abnormal
visible dorsolaterally but with rigid internal (probably damaged during postembryonic de-
sclerotized ridge instead, furnished with 8 large velopment). Outer apical seta without defor-
sensilla dorsally (Fig. 11); additionally orna- mities, also with breaking plane, about 2.8 times
mented with 6 dorsal, 2 ventral, and 2 lateral as long as inner apical seta, smooth or pinnate
sensilla near posterior margin; hyaline fringe along very short section (Fig. 1).
sharply serrated both ventrally and dorsally. Antennula (Fig. 2) 8-segmented, very short,
Female genital complex with single large approximately 1.5 times as long as rostrum,
copulatory pore and without epicopulatory bulb; with slender aesthetasc on eighth segment and
copulatory duct short, cylindrical, rigidly scler- longer and stouter aesthetasc on fourth segment,
otized; 2 small seminal receptacles reniform, which exceeding in length the last six segments
placed inside single large genital aperture; together; setal formula as follows: 1.9.4.4.1.
aperture covered by fused and reduced sixth 4.4.7. No seta with articulating base or breaking
legs (Fig. 7). Third urosomal somite ornamented plane. Seta on first segment, 1 seta on second, 2
with 4 dorsal, 4 ventral, and 2 lateral sensilla and on third, 1 on fourth, 2 on sixth, and 1 on seventh
with short ventral row of large spinules near segment pinnate; all other setae smooth. Length
posterior margin; fringe sharply serrated both ratios of antennular segments, from proximal end
dorsally and ventrally, although ventral teeth and measuring medially, 1 : 0.6 : 0.5 : 0.3 : 0.3 :
larger. Preanal somite without any sensilla or 0.3 : 0.3 : 0.8. First segment ornamented with
spinules; hind margin clearly bulging posteriorly transverse row of spinules; other segments
in dorsal region, forming very sharply serrated without any ornamentation visible.
pseudoperculum (Fig. 11); hyaline fringe frilled Antenna (Fig. 3) comprising coxa, allobasis,
dorsally and serrated both dorsally and ventrally. 1-segmented endopod, and 3-segmented exopod.
Anal somite without anal operculum; orna- Coxa very short, about twice as wide as long,
mented with 2 large sensilla and 2 cuticular unornamented. Basis and proximal endopodal
pores dorsally, as well as with transverse row of segment fused forming allobasis, without origi-
large spinules on posterior margin ventrally and nal segmentation marked by any surface suture,
laterally (Figs. 7, 11). Anal sinus smooth. armed with only 1 strong pinnate seta on anterior
Caudal rami (Figs. 7, 10, 11) cylindrical in margin; ornamented with transverse row of very
dorsal or ventral view, but slightly conical in long spinules between exopod and anterior
lateral view, with posterior margins terminating margin. Single free endopodal segment with 2
ventrally in narrow acuminate lappets (between surface frills subdistally; lateral armature con-
middle and outer apical seta), divergent, with sisting of 2 spines and 1 small seta; apical
space between them being about third of ramus armature consisting of 1 slender seta (which
width, and about 1.7 times as long as greatest bipinnate at distal end), 1 short spine, 4 strong
 KARANOVIC AND RANGA REDDY: NEOMISCEGENUS, N. GEN., FROM INDIA 255

geniculate setae, longest one bearing very long Syncoxa ovoid, with 3 endites; proximal and
spinules around geniculation and fused basally to middle endite each armed with 2 short subequal
another, smooth slender seta; ornamentation setae, distal endite armed with 3 subequal setae.
consisting of longitudinal row of very large Basis drawn out into strong claw, armed with 1
spinules along anterior margin, diagonal row of strong and 2 slender setae at base. Each
large spinules between lateral and apical arma- endopodal segment armed with 2 slender setae.
ture elements, and diagonal row of small spinules Ornamentation on maxilla consists of few small
along subapical frill. First exopodal segment spinules on outer margin of syncoxa.
unornamented and about twice as long as wide, Maxilliped (Fig. 14) prehensile, 4-segmented,
armed with 1 pinnate subapical seta on anterior composed of coxa, basis, and 2-segmented endo-
distal corner; second segment minute, unarmed pod. Coxa reduced, unornamented. Basis orna-
and unornamented (on right antenna even partly mented with arched row of large spinules and
fused to first segment (Fig. 6)), about twice as armed with 2 pinnate setae at distal inner corner.
wide as long; third exopodal segment conical, First endopodal segment about 2.5 times as long
ornamented with transverse row of spinules at 2/ as wide, ornamented with 2 longitudinal rows of
3 of its length, armed with 1 bipinnate apical seta spinules and armed with 1 lateral smooth and 1
and 1 unipinnate lateral seta, which inserted at 1/ subapical unipinnate seta. Second endopodal seg-
3 of segment’s anterior margin. ment small, armed with 1 claw-like apical spine
Labrum (Fig. 4) large, rigidly sclerotized, and 2 slender and much shorter subapical setae
with trapeziform free part and ornamented with and ornamented with single subapical spinule.
1 short row of spinules near each posterior outer All swimming legs with 3-segmented exopods
corner; cutting edge slightly convex, smooth. and endopods (Figs. 16, 17, 18, 19). Swimming
Mandibula (Figs. 8, 9) with broad cutting legs armature formula as follows (legend: inner/
edge of coxa, armed with many pointed teeth outer spine or seta; inner/terminal/outer):
and with 1 unipinnate seta. Basis elongate,
Exopod Endopod
about 2.3 times as long as wide, armed with
3 slender pinnate setae along inner margin. Segments 1 2 3 1 2 3
Endopod 1-segmented, about 1.7 times as long First leg 0/1 0/1 0/2/2 1/0 1/0 1/1/1
Second leg 0/1 1/1 0/2/3 1/0 1/0 1/2/1
as wide, armed with 2 lateral and 4 apical setae. Third leg 0/1 1/1 1/2/3 1/0 1/0 2/2/1
Exopod also 1-segmented but much smaller Fourth leg 0/1 1/1 2/2/3 1/0 1/0 1/2/1
than endopod, about 1.9 times as long as wide,
armed with 2 lateral and 2 apical smooth setae. Intercoxal sclerites of all swimming legs with
Both lateral setae on endopod inserted at concave distal margins; sclerite of first leg small
middle, while on exopod 1 lateral seta inserted and smooth (Fig. 16); sclerite of second leg
at 1/3 and other at 2/3 of segment length. armed with 2 arched rows of large spinules at
Maxillula (Fig. 12) with large praecoxa, distal part (Fig. 17); sclerite of third leg smooth
arthrite of which highly mobile, armed apically but with pointed outer distal corners (Fig. 18);
with 4 smooth strong spines, laterally with 2 and sclerite of fourth leg smooth and with
smooth slender anterior surface setae, dorsally rounded outer distal corners (Fig. 19). Prae-
with 2 short pinnate setae; ornamented dorsally coxae of all legs with transverse row of
with 2 long spinules. Coxa small, armed with spinules, except fourth leg smooth. Coxae orna-
single smooth strong seta on inner margin, not mented with at least 2 rows of large spinules, un-
reaching midlength of basis. Basis slightly armed. Basis of first swimming leg armed with
longer than arthrite of praecoxa, furnished with 1 strong spine on inner distal corner (reaching
3 setae apically (on inner margin) and with 2 2/3 of first endopodal segment) and 1 moderate
setae laterally; ornamented with longitudinal spine on outer margin; ornamented with 1 cres-
row of small spinules. Endopod 1-segmented, centic row of large spinules at base of inner
small, about twice as long as wide, armed with 1 spine, 1 similar row of spinules on distal mar-
lateral and 3 apical setae, median longest. gin, between endopod and exopod, and with
Exopod also 1-segmented, about 0.7 times as short row of minute spinules on inner margin.
long as endopod and 1.7 times as long as wide; Basis of second leg with spine on outer margin;
armed with 2 plumose apical setae, outer seta those of third and fourth legs with strong seta.
being about 1.7 times as long as inner one. Inner distal corners of basis of second, third and
Maxilla (Fig. 13) composed of syncoxa, fourth legs each produced distally as sharp,
basis, and very small 2-segmented endopod. spiniform process; that of second leg being
256 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 24, NO. 2, 2004

longest. Endopods of first, second, and third ventral acuminate lappets and armature. Inner
legs slightly longer than exopods, endopod of apical seta smooth and slightly shorter than
fourth leg considerably shorter than exopod. lateral one; ventral seta inserted at 3/4 of ramus
First endopodal segment of first swimming leg length; middle apical seta about twice as long as
reaching middle of second exopodal segment outer apical one, and both setae with breaking
(Fig. 16), about 2.4 times as long as wide. planes.
Second endopodal segments of second, third, Antennula (Fig. 23) 9-segmented (although
and fourth legs each with sharply produced distal part of third segment distinct lobe bearing
outer distal corner. All exopods and endopods 1 apical seta) not strongly geniculate, with
ornamented with spinules on outer margins, and geniculation visible between sixth and seventh
exopods also ornamented with hairs on inner segments, somewhat longer than in female.
margins. All armature elements very strong, Aesthetascs present on fourth and ninth seg-
except inner subapical seta on third endopodal ments; first aesthetasc much longer and stouter.
segment of first leg. Setal formula as follows: 1.11.8.6.2.3.0.4.4.
Fifth leg (Fig. 15) biramous. Baseoendopod Seta on first segment and 1 seta on third
with outer basal seta smooth, arising from segment pinnate; all other setae smooth. Only
relatively short setophore. Endopodal lobe sub- 1 seta on ninth segment articulating on basal
trapeziform, extending to 2/3 of exopod length, part and no setae with breaking planes; 2 setae
ornamented with few spinules on inner margin on sixth segment transformed into spiniform
and armed with 5 armature elements; 2 inner armature elements. Seventh segment with char-
elements very stout, spiniform, unipinnate and acteristic cuticular structure on grasping margin.
longer than baseoendopod; 3 outer elements Antenna, labrum, mandibula, maxillula, max-
also very strong, but bipinnate and shorter than illa, maxilliped, endopod, and exopod of first
baseoendopod; length ratios of 5 endopodal swimming leg, exopod of second swimming leg
setae (from inner side) approximately 1 : 1.1 : (Fig. 24), and fourth swimming leg similar to
0.7 : 0.8 : 0.5. Exopod subquadrangular, about female.
1.6 times as long as maximum width, un- First swimming leg (Fig. 28) with modified
ornamented, and armed with 5 armature ele- basis, inner margin of which rigidly sclerotized
ments; 2 inner setae subequal, very strong, and produced distally into blunt spiniform
spiniform and bipinnate; middle seta slender, process, at base of inner spine. Inner spine on
smooth and longest one; 2 outer setae subequal, basis somewhat smaller than in female, without
very short, curved and unipinnate; length ratio spinules at its base, about 2.4 times as long as
of 5 exopodal setae (from inner side) 1 : 1.3 : spiniform process.
1.7 : 0.4 : 0.4. Second swimming leg (Fig. 26) with trans-
Sixth legs (Fig. 7) fused, indistinct, very small formed endopodal second and third segments.
cuticular plates, covering single large gonopore, Second segment with part of inner margin
armed with 2 setae each; inner seta 1.8 times as protruding into smooth rounded lobe; inner seta
long as outer one, slender and smooth and shorter than in female. Third segment completely
reaching posterior end of seminal receptacles; modified; only inner seta normally formed, stout,
outer seta stouter than inner one and plumose. bipinnate, and very long. Ancestral inner apical
Male (allotype). Body length, excluding seta smooth and very slender, shorter than inner
caudal setae, 0.327 mm. Habitus, ornamentation seta. Ancestral outer apical seta transformed into
of prosomal somites, rostrum, dorsal ornamen- strong smooth serpentine element; its distal end
tation of urosomal somites, pseudoperculum swollen and with 2 characteristic tips (1 apically
(Fig. 20), color and nauplius eye similar to and 1 on outer margin). Ancestral outer spine
female. Genital somite about 2.6 times as wide completely fused to somite, transformed into
as long. Single large, completely formed and very strong and smooth thorn, which somewhat
longitudinally placed spermatophore (Fig. 22) shorter than ancestral outer apical element and as
visible inside first 2 urosomal somites. All long as ancestral inner apical seta; with small
abdominal somites ventrally with posterior rows smooth protrusion at inner basal margin. As
of spinules; these rows interrupted only on pre- a result of these transformations, third segment
anal somite, and first 3 somites with additional appearing cleft medially. Exopod (Fig. 24)
pair of sensilla in each row (Fig. 21). similar to female.
Caudal rami (Fig. 21) much shorter and Third swimming leg (Fig. 25) with sexually
less ornamented than in female, but with similar dimorphic pore on third exopodal segment, at
 KARANOVIC AND RANGA REDDY: NEOMISCEGENUS, N. GEN., FROM INDIA 257

level of second outer spine, but without hyaline genera, together with Amphiascoides Nicholls,
tubular extension. 1941 [syn. Amphiascella Lang, 1944], are very
Fifth legs (Fig. 27) with basally fused base- close to the Indian freshwater representative
oendopods and without ornamentation. Endopo- insomuch that they share the same female
dal lobe small, trapeziform, extending to swimming legs armature formula, as well as
midlength of exopod, armed with 2 very strong the fifth leg armature formula in both sexes.
apical spines; inner spine about 1.1 times as long Unfortunately, the Indian species cannot be as-
as outer one. Exopod about as long as its signed to any of these four genera (or any other
maximum width, pentagonal; armed with 5 genus of Diosaccidae) without significantly
elements, as in female; length ratios of these extending their diagnoses. As that would cause
elements (from inner side) 1 : 1.5 : 1.8 : 0.3 : 0.4. systematic confusion in many other genera, we
Sixth leg (Fig. 29) distinct, broad and short, prefer to erect a new genus to accommodate it.
cuticular plate, without ornamentation and The genus Amphiascoides contains today 20
armed with 3 elements: smooth innermost spine, valid species, fewer than when it was originally
slender middle seta (this seta slightly shorter established by Nicholls (1941). The reason for
than spine and also smooth) and minute that is its junior synonym, the genus Amphias-
outermost seta (about twice as short as spine). cella, described by Lang (1944), which di-
agnostic concept was much narrower and has
Variability.—Exopod of antenna may have all been accepted consequently. That is why all
three segments distinct (Fig. 3), or last two generic comparisons should refer to Lang’s
segments can be partly fused (Fig. 6). Un- (1948, 1965) perception of this group of
fortunately, only one male and one female were Diosaccidae species, of course, with Nicholls’
collected and studied. (1941) name having priority. Although a rela-
Etymology.—The species name is taken from the tively close relationship between Amphias-
name of the Republic of India, where the material coides and Neomiscegenus, n. gen., is evident,
was collected, i.e., as an adjective agreeing in the former differs from the latter by some very
gender with the masculine generic name. important morphological features: the first
endopodal segment of first leg is always longer
than the entire exopod; the caudal rami are
DISCUSSION wider than long, and the proximal lateral seta is
Compared with many other harpacticoid taxa, not moved completely ventrally; the third
the subdivisions of Diosaccidae Sars, 1906, are exopodal segment of the antenna bears more
more or less clear, despite the fact that this large than one apical seta; the exopod of the
family has been treated independently by mandibula is 3-segmented; the basis of the
Nicholls (1941) and Lang (1948). Lang was maxilliped is armed with 3 setae; the female
apparently unaware of Nicholls’ paper when he antennula is much longer (at least twice, but
wrote his monograph. Although Lang’s system usually more than three times, as long as the
is far superior, and consequently has been rostrum); the epicopulatory bulb on the female
adopted by all modern taxonomists, Nicholls genital somite is present; and the endopod of the
introduced new generic names that cannot be second leg in the male is much more simplified.
neglected for priority reasons. Lang’s second With 21 species currently recognized, the
overview of this family (Lang, 1965) certainly genus Paramphiascella appears to be as large as
greatly contributed to the clarity of the generic Amphiascoides, but the former probably con-
diagnoses, but some genera have not been tains several additional unrecognized syno-
properly assigned until recently (see Huys, nyms. Regarding this genus, Lang (1965: 321)
1990). Lang’s (1948) view of the phylogeny of stated: ‘‘the differences between many of the
Diosaccidae has not been challenged so far, species are so small that I cannot give a key to
although it was supplemented by Wells et al. them.’’ Whatever is the exact number of species
(1982) for the Amphiascus-related genera. Our in Paramphiascella, they form quite a compact
new species clearly belongs to that evolutionary entity and can be distinguished from Neo-
line and, in fact, in Lang’s (1965) key it would miscegenus by the following six characteristics:
outkey as Paramphiascella Lang, 1948, just like the first endopodal segment of the first leg is
the genera Paramphiascoides Wells, 1967, and always longer than at least the first two
Miscegenus Wells, Hicks, and Coull, 1982 (see exopodal segments combined; the third exopo-
Wells, 1967; Wells et al., 1982). These three dal segment of the antenna bears more than one
258 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 24, NO. 2, 2004

apical seta; the exopod of the mandibula is 2- tory bulb is present; the exopod of the maxillula
segmented; the female antennula is much bears three setae; and, most importantly, the
longer, when compared with the cephalothorax endopod of the second leg in the male is quite
or rostrum, and slender; the epicopulatory bulb different from that in Neomiscegenus.
on the female genital double-somite is present; As its name suggests, Neomiscegenus is
and the endopod of the second leg in the male morphologically most similar to the genus
ends like slender pincers, without any trace of Miscegenus, which is also known from a single
the third segment itself. Obviously, Param- species. This species, Miscegenus hereatunga
phiascella is much closer to Amphiascoides than Wells, Hicks, and Coull, 1982, was described
to Neomiscegenus, and the only strong generic from several estuaries and bays from New
character that separates them is the appearance Zealand by Wells et al. (1982). It shares with
of the sexually dimorphic endopod of the Neomiscegenus indicus a number of important
second leg in the male (Lang 1965: 320). characters, including the short antennulae;
However, this seems to be enough, and we fusiform habitus; similar caudal rami (with the
completely agree with Mu and Gee (2000: 129) ancestral proximal lateral seta moved ventrally,
that: ‘‘The secondary sexually dimorphic char- and a ventral lappet between the outer and
acters may assume greater significance in future middle apical setae); the same armature formula
revisions of the genera of Diosaccidae . . .’’ It of all the swimming legs and the fifth legs;
should be mentioned here that some of the similar maxilliped, the endopod of the second
characters that we are using to distinguish leg in the male with similar basic structure; and
Neomiscegenus from Paramphiascella (as well even extremely similar ornamentation of uro-
as from Amphiascoides) are not known in all somal somites, in both female and male.
species of the latter genus, because of the Because it would be impossible to consider all
incomplete descriptions of many species, but these morphological similarities as convergen-
unfortunately, that is the present situation in the cies, the two genera must have a close phylo-
majority of copepod genera. It is interesting to genetic relationship. Nevertheless, a few very
mention that one of the Paramphiascella important characters preclude us from assigning
species, P. aquaedulcis Dussart, 1984, was the new Indian species to the genus Miscegenus.
described from a similar habitat as Neomisce- The most important one is certainly the absence
genus indicus, n. sp., after the seasonal flooding of an epicopulatory bulb in the Indian repre-
of Rio Portuguesa in Venezuela (Dussart, 1984). sentative (Fig. 7), because Miscegenus pos-
Probably this evolutionary line of Diosaccidae sesses a large bulb on the female genital
genera is more successful in colonizing the segment, which is much more like that in the
freshwater subterranean habitats than we pres- genus Schizopera Sars, 1905, than in Amphias-
ently know, and further investigations of this coides, Paramphiascella, or Metamphiascopsis
habitat in former parts of Gondwana may reveal Lang, 1948 (see Ohtsuka and Iwasaki, 1998).
many new members. Although the function of this female sexual
The genus Paramphiascoides is so far mono- structure is unknown, Huys and Boxshall
specific. It was described from the littoral of (1991) considered it as a ‘‘novel structure’’ in
Inhaca Island, Mozambique, by Wells (1967). Harpacticoida. That all 70 species and sub-
Its sexually dimorphic endopod of the second species of the genus Schizopera have this
leg in the male ends in slender pincers, just as in structure, which can even be used sometimes
the genus Paramphiascella, but it has quite as one of the additional characters in distin-
a different appearance. None of the two guishing closely related species (Karanovic,
branches of this peculiar organ is movable, 2004), is proven. It is hard to imagine that the
and also the inner setae are not so closely set as epicopulatory bulb arose independently a num-
in Paramphiascella. The first swimming leg of ber of times within the Diosaccidae. Because the
the genus Paramphiascoides is very similar to condition in the female genital field is not one of
that of Neomiscegenus, and the female anten- the common stygomorphies (character states,
nulae are also very short. However, Param- usually reductions, caused by a subterranean
phiascoides can be easily distinguished from the aquatic life) in copepods, we consider it as
new taxon by the following characters: the a good generic character. The endopod of the
distalmost exopodal segment of the antenna second leg in the male is similar in both
bears more than one apical seta; the exopod of Miscegenus and Neomiscegenus, but it has
the mandibula is 3-segmented; the epicopula- several very important differences: the third
 KARANOVIC AND RANGA REDDY: NEOMISCEGENUS, N. GEN., FROM INDIA 259

segment with only one lateral seta in Neo- (1948) phylogenetic tree made by Wells et al.
miscegenus (versus two in Miscegenus); the (1982) are not acceptable, because they are
apical seta on it shorter than lateral (longer in focused on one character only and, as a result,
Miscegenus); and the ancestral outer apical seta the genus Miscegenus is misplaced. We think
transformed into a serpentine element with two that the genera Amphiascoides, Paramphias-
characteristic tips in Neomiscegenus (almost cella, Paramphiascoides, Miscegenus, and Neo-
straight in Miscegenus and without any tips). miscegenus have the same common ancestor.
This scale of differences is not known within Within this line, during their evolutionary
any of the Diosaccidae genera, so we consider it history, the genera Miscegenus and Neomisce-
as a generic feature as well. However, tips on genus have separated relatively early from the
the second leg endopod of Neomiscegenus are other branch, which evolved later into the
not an autapomorphy, but rather a plesiomorphy, genera Amphiascoides, Paramphiascella, and
because they can be found in the genera Paramphiascoides.
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