Three new species of Parastenocarididae (Crustacea, Copepoda, Harpacticoida) from India

Zootaxa 3821 (5): 501–537 www.mapress.com /zootaxa / Copyright © 2014 Magnolia Press

Article

ISSN 1175-5326 (print edition)

ZOOTAXA

ISSN 1175-5334 (online edition)

http://dx.doi.org/10.11646/zootaxa.3821.5.1 http://zoobank.org/urn:lsid:zoobank.org:pub:E3CDE3FE-22C3-431B-BD9D-D254721E6C50

Three new species of Parastenocarididae (Crustacea, Copepoda, Harpacticoida) from India VENKATESWARA RAO TOTAKURA1, YENUMULA RANGA REDDY1,2 & SHABUDDIN SHAIK1 1

Department of Zoology, Acharya Nagarjuna University, Nagarjunanagar 522 510, India Corresponding author. E-mail: [email protected]

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Abstract To date, only 16 species of the very diverse family Parastenocarididae have been reported from the Indian subcontinent. This paper gives an illustrated description of three new species, viz. Parastenocaris edakkal n. sp., Proserpinicaris corgosinhoi n. sp., and Proserpinicaris karanovici n. sp., and discusses their position in the respective genera. While the first two species were found sympatrically in the Edakkal Cave in Kerala State of southwestern India, the third one was collected from a farm bore in the riparian zone of the River Krishna in Andhra Pradesh State of southeastern India. The Palaearctic genus Proserpinicaris Jakobi, 1972 sensu Karanovic, Cho & Lee, 2012, is being reported for the first time from India. Parastenocaris edakkal n. sp. belongs to the brevipes-group of the genus Parastenocaris Kessler, 1913 sensu Lang, 1948 et Reid, 1995. It is chiefly characterized by the male leg 4 basal chitinous complex consisting of one large sclerotized plate together with two small accessory lobes, and one strong, hook-like spine at the inner distal corner, and the endopod is membranous and ventricose in outline, with bulbous proximal part drawn out distally into biserrulate, pointed structure. Proserpinicaris corgosinhoi n. sp. can be easily separated from its congeners, inter alia, by the short caudal rami (c.1.2–1.4 times as long as wide), bearing 2 unequal lateral setae (I, III) inserted proximally; male leg 3 proximal segment is stumpy and has one prominent tubular pore on proximal anterior surface; and leg 4 endopod is short, membranous and somewhat conical, and the hyaline structure is relatively large, foliaceous and lies rather close to endopod. Proserpinicaris karanovici n. sp. has its own unique set of characters, of which the following are noteworthy: the caudal rami in both sexes are about 2.6 times as long as wide, gradually tapering, with only 2 lateral setae inserted in proximal half; and the male leg 4 endopod is nearly as long as first exopodal segment, membranous, with lateral margins fringed with tiny spinules, and the hyaline structure is short, leaf-like and occurring close to, and overlapping, the endopod. Key words: Parastenocaris edakkal n. sp., Proserpinicaris corgosinhoi n. sp. Proserpinicaris karanovici n. sp., stygofauna, taxonomy

Introduction The family Parastenocarididae is presently known by only 16 species on the Indian subcontinent, as against the world tally of over 290 described species and subspecies in 30 genera (Gaviria-Melo et al. 2013). Of these 16 species, all but the Indian Kinnecaris godavari Ranga Reddy & Schminke, 2009 and Siolicaris sandhya (Ranga Reddy, 2001) belong to the nominotypical genus Parastenocaris Kessler, 1913. While six of the 13 remaining species of Parastenocaris are from Sri Lanka (Enckell 1970), viz. Parastenocaris irenae Enckell, 1970, Parastenocaris noodti Enckell, 1970, Parastenocaris brincki Enckell, 1970, Parastenocaris singhalensis Enckell, 1970, Parastenocaris lanceolata Enckell, 1970, and Parastenocaris curvispinus Enckell, 1970, eight species including the Sri Lankan P. curvispinus, are distributed in India, viz. Parastenocaris gayatri Ranga Reddy, 2001, Parastenocaris savita Ranga Reddy, 2001, Parastenocaris mahanadi Ranga Reddy & Defaye, 2007, Parastenocaris muvattupuzha Ranga Reddy & Defaye, 2009, Parastenocaris kotumsarensis Ranga Reddy & Defaye, 2009, Parastenocaris sutlej Ranga Reddy, 2011, Parastenocaris gundlakamma Ranga Reddy, 2011, and Parastenocaris tirupatiensis Ranga Reddy, 2012. Incidentally, P. curvispinus is most common in peninsular India (Ranga Reddy & Defaye 2007). All the hitherto described Indian species are hyporheic except for P. kotumsarensis

Accepted by T. Karanovic: 20 May 2014; published: 25 Jun. 2014

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and P. tirupatiensis, which were collected from a karstic cave pool and pumped-out water of a farm bore, respectively (see Ranga Reddy 2001, 2011a, 2012; Ranga Reddy & Defaye 2007, 2009). During our ongoing investigations on the Indian stygofauna, we have encountered seven more new taxa of Parastenocarididae. This paper gives an illustrated description of three new species, of which two species, viz. Parastenocaris edakkal n. sp. and Proserpinicaris corgosinhoi n. sp., both co-existing in the interstitial waters of the Edakkal Cave in the Wayanad District of Kerala State, southwestern India, and a new phreatic Proserpinicaris karanovici n. sp. was found in a farm bore in the riparian zone of the River Krishna near Vijayawada city, southeastern India (Fig. 1a). While P. edakkal n. sp. belongs to the brevipes-group, as redefined by Reid (1995), of the genus Parastenocaris Kessler, 1913 of the subfamily Parastenocaridinae Chappuis, 1940, the other two new species belong to the subfamily Fontinalicaridinae Schminke, 2010. This is the first record of the Palearctic genus Proserpinicaris Jakobi, 1972, as revised by Karanovic et al. (2012), from the Indian subcontinent.

Material and methods Specimens were collected with two methods: 1. The specimens of Parastenocaris edakkal n. sp. and Proserpinicaris corgosinhoi n. sp. were collected from the interstitial water of core samples taken from fine sand mixed with silt and organic debris in the Edakkal cave, Kerala, southwestern India (see Type locality and Fig. 1a). A rigid PVC tube (70 cm length, 4 cm diameter) was used to extract cores from the sediment surface to a depth of 10–20 cm in a small, shallow cave pool atop a hill. The samples were pooled in a bucket, filled with spring-originated, flowing water near the sampling spot (examination of a plankton sample from the flowing water showed no trace of any animals) and stirred vigorously. The supernatant was filtered through a bolting-silk plankton net (70 µm mesh size) and the filtrate fixed in 5% formaldehyde. The specimens, which were obviously interstitial, were sorted into 70% alcohol and later transferred into glycerol. 2. The specimens of Proserpinicaris karanovici n. sp. were collected by directly filtering the groundwater pumped from a farm bore (depth c. 10 m) with a plankton net for 20–30 minutes (see Fig. 1b for the method of collection). Specimens were dissected in glycerol under a binocular stereo zoom microscope at a magnification of 90×. Drawings were made with the aid of a drawing tube mounted on a Leica DM 2500 Trinocular Research Microscope equipped with UCA condenser, IC objective prism and 1–2× magnification changer. Permanent preparations were mounted in glycerol and sealed with wax and Araldite. All the type material was deposited in the Muséum national d’Histoire naturelle (MNHN), Paris.

Systematics Subphylum Crustacea Brünich, 1772 Class Maxillopoda Dahl, 1956 Subclass Copepoda H. Milne Edwards, 1840 Order Harpacticoida G.O. Sars, 1903 Family Parastenocarididae Chappuis, 1940 Subfamily Parastenocaridinae Chappuis, 1940 Genus Parastenocaris Kessler, 1913

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FIGURE 1. (a) Map showing the distribution of the Indian parastenocaridid species including Parastenocaris edakkal n. sp., Proserpinicaris corgosinhoi n. sp., and Proserpinicaris karanovici n. sp.; (b) sampling of a farm bore.

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Parastenocaris edakkal n. sp. (Figs 2–8) Type locality. Edakkal cave (11°37′54.0″N, 76°14′14.9″E, elevation 1233 m, water temperature 30ºC, pH 7.0) in Wayanad District of Kerala State in the Western Ghats, which constitute one of the key biodiversity hotspot areas of India (Fig. 1a). This Cave is not technically a cave, but rather a cleft or rift of about 20 m long, 6.7 m wide and 9.1 m deep fissure formed by a piece of rock (probably Kondalite) splitting away from the main body. On one side of the cleft is a huge rock covering the cleft to form the cave roof. Pictorial Stone Age carvings of human and animal figures and tools used by Neolithic man are seen inside the cave. These carvings are said to be as old as 5000-6000 B. C., suggesting the prehistoric civilization or settlement in this region (see http://www.edakkal.com/ index.htm). A small stream, probably originating from a perennial underground spring, flows through the length of the cave over fine sand deposit mixed with organic debris. There are no drippings from the rocky roof. Type material examined. Holotype male (MNHN–IU–2013–11244) and allotype female (MNHN–IU–2013–11245) dissected on 3 slides each and 10 paratypes: 1 male (MNHN–IU–2013–11246) and 1 female (MNHN–IU–2013–11247) whole-mounted on 1 slide each, 1 male and 7 females in alcohol in a vial (MNHN–IU–2013–11248). 09 May 2008, Coll. V. R. Totakura and Y. Ranga Reddy. Description of adult male. Total body length, measured from tip of rostrum to posterior margin of caudal rami (excluding caudal setae), 368 μm. Body heavily chitinized and pitted. Naupliar eye absent. Habitus (Fig. 2a, b) cylindrical and slender, without any podoplean demarcation between prosome and urosome; prosome/urosome ratio about 0.7 in dorsal view; greatest width in dorsal view falling in distal half of cephalothorax. Body length/ width ratio about 7.8. Free pedigerous somites without any lateral or dorsal expansions. All somites connected by well developed arthrodial membranes. Hyaline fringes of all somites smooth, very narrow and hard to distinguish from arthrodial membranes. Integument ornamented only with sensilla, cuticular pores, pits (no spinules), and also with sub-spherical, dorsal double-window on cephalothorax, and elliptical, dorsal simple cuticular window each on genital somite and next 3 somites, window on genital somite smallest. Pleural areas of cephalothorax moderately developed; cephalic appendages and coxae of legs 1–4 only partly exposed in lateral view. Rostrum (Fig. 2a, b) small, membranous, not demarcated at base, ornamented with 2 large dorso-lateral sensilla. Cephalothorax (Fig. 2a, b): subquadrate, about 1.2 times as long as wide in dorsal view, 1.3 times in lateral view, and representing 16% of total body length. Surface of cephalic shield ornamented with 4 pairs of large sensilla on anterior half as well as around cuticular double-window; window surface completely smooth, no cuticular pores discernible; second and third pedigerous somites as wide as posterior part of cephalothorax in dorsal view, with 3 and 2 pairs of sensilla (dorsal and lateral), respectively; third somite slightly shorter than second segment with 3 pairs of sensilla. Fourth pedigerous somite longer than preceding somite, with 3 pairs of sensilla (dorsal, lateral). Urosome (Figs. 2a, b, 3a): first urosomite about as wide as fourth prosomite, but slightly shorter, ornamented with 3 pairs of sensilla dorsally (Fig. 2b) and also, 1 sensillum beside leg 5 ventrally (Fig. 3a), 1 pair of lobes in proximal-half ventrally (Fig. 3a). Genital somite slightly wider than first urosomite, with small, oval dorsal cuticular window in anterior half, also with 2 pairs of sensilla (dorsal, lateral) and 1 pair of sensilla (ventral) and 1 sensillum on either side of sixth legs. Third and fourth urosomites about as long as first urosomite but slightly narrower, with wider dorsal cuticular window each, and with 2 pairs of large posterior sensilla on distal half and 1 pair of sensilla disto-ventrally. Preanal somite narrower and slightly longer than fourth urosomite, with largest dorsal cuticular window, and without any surface ornamentation. Anal somite about 1.2 times as long as and slightly narrower than preanal somite, 1.2 times as long as its own width; ornamented with 1 pair of large dorsal sensilla and 1 antero-lateral cuticular pore proximally (Fig. 2a). A single large, longitudinally placed spermatophore discernible through cuticle of fourth and fifth segments, about 3.1 times as long as wide, kidneyshaped, with narrow curved neck. Anal operculum well developed, almost reaching posterior end of distal margin, with smooth and nearly straight distal margin, representing 73.3% of somite's width, unornamented on outer surface, transverse row of ventral spinules discernible through transparent operculum. Anal sinus wide open. Caudal rami (Figs. 2a, b, 3a): divergent, cylindrical, gradually tapering posteriorly, about 3.3 times as long as greatest width in lateral view (Fig. 2a), 3.2 times in dorsal view (Fig. 2b), and about 0.6 times as long as anal somite, ornamented with 1 row of tiny spinules on ventral surface disto-laterally and with 1 distolateral pore, and armed with full complement of 7 setae (3 lateral, 1 dorsal, 1 subapical and 2 apical). Setae I–III thin, unequal, located as a group at about the middle of ramus. Dorsal seta (VII) slender and unipinnate, biarticulate basally, inserted closer to inner margin and opposite the level of setae I–III, about 0.6 times as long as caudal ramus. Inner apical seta (VI) smooth, proximally swollen, inserted close to ventral margin, about 0.6 times as long as ramus.

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Middle apical seta (V) strongest, without breaking plane, unipinnate, with slightly curved tip. Outer subapical seta (IV) strong basally, also without breaking plane and unipinnate, about as long as ramus, inserted close to dorsal surface and directed laterally.

FIGURE 2. Parastenocaris edakkal n. sp., paratype male: (a) habitus, lateral view; (b) habitus, dorsal view. NEW INDIAN SPECIES OF PARASTENOCARIDIDAE

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FIGURE 3. Parastenocaris edakkal n. sp., (a) holotype male, urosome, ventral view; (b) allotype female, urosome, dorsal view.

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FIGURE 4. Parastenocaris edakkal n. sp., paratype female (a, c), allotype female (b): (a) habitus, lateral view; (b) urosome, ventral view; (c) leg 5, latero-ventral view.

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FIGURE 5. Parastenocaris edakkal n. sp., holotype male: (a) antennule, dorsal view; (b) antenna, lateral view; (c) labrum, ventral view; (d) mandible, lateral view; (e) maxillule, lateral view; (f) maxilla, lateral view; (g) maxilliped, lateral view.

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Antennule (Fig. 5a): 1.3 times longer than cephalothorax, slender, 8-segmented, and prehensile, ‘pocket-knife type’, digeniculate, proximal geniculation between third and fourth segments, and distal geniculation between sixth and seventh segments. First segment very short, ornamented with 1 row of spinules. Second one longest. Segments 4–6 moderately dilated, segment 5 without proximal spinous process on anterior surface; elongate aesthetasc with pointed tip, overreaching midlength of aesthetasc on ultimate segment, fused basally to 1 simple seta; shorter and slenderer apical aesthetasc on eighth segment, fused basally with 2 setae (acrotheck). Setal formula: 0.5.3.1.2+aes.0.1.8+aes. All setae slender, smooth except proximalmost seta on second segment, which is unipinnate, with long setules along outer margin. Length ratios of antennular segments, from proximal to distal end along caudal margin 1:2.2:0.8:0.3:1.7:0.5:0.7:1.1. Antenna (Fig. 5b): relatively stout, composed of coxa, allobasis, 1-segmented endopod, and 1-segmented exopod. Coxa very short, ornamented with row of short spinule, and unarmed. Allobasis about 2.4 times as long as maximum width, unarmed, ornamented with 1 oblique row of spinules near mid-inner margin. Exopod small, cylindrical, about 2.3 times as long as wide, unornamented, armed with apical unipinnate seta, which is 2.8 times as long as segment. Endopod 0.6 times as long as allobasis and about twice as long as wide, surface frill occurring distally, ornamented with 1 large and 1 small arched spinular rows on inner margin, armed with 2 short bipinnate, unequal spines laterally and with 5 strong elements apically (2 subequal spines, 2 subequal geniculate setae and 1 unipinnate transformed seta). Labrum (Fig. 5c): large, triangular in lateral view (Fig. 4a) and sub-ovate in ventral view, with narrow, concave, cutting edge finely denticulate, with short spinous projection on either side; also, ornamented with 1 row of arched spinules on ventral surface close to cutting edge. Mandible (Fig. 5d): cutting edge narrow on elongate coxa, armed with 2 complex teeth ventrally, 1 unipinnate seta dorsally, and several smaller teeth. Palp 1-segmented, cylindrical, about 3 times as long as wide, unornamented, and armed apically with 2 smooth, unequal apical setae. Maxillule (Fig. 5e): with relatively large praecoxa, arthrite rectangular, about 3.3 times as long as wide in lateral view, armed with lateral stout curved seta, and 3 apical claws of which middle one with serrulate outer margin. Coxal endite armed with 2 smooth, unequal setae apically. Basis 1.3 times longer than coxal endite, armed with 1 strong and 2 slender smooth apical setae. Maxilla (Fig. 5f): composed of syncoxa, basis, and 1-segmented endopod. Syncoxa with 2 endites, proximal one short, armed with 2 smooth unequal setae apically, distal endite armed with 1 spiniform and 1 smooth setae apically. Allobasis prolonged into strong, unipinnate claw, without seta at base. Endopod represented by small segment, armed with 2 smooth apical setae. Maxilliped (Fig. 5g): with short and relatively strong syncoxa, unornamented and unarmed; basis slender, 2.5 times as long as wide and 2.1 times as long as syncoxa, unornamented and unarmed; endopod small with unipinnate claw, 0.6 times as long as basis. Legs 1–4 (Figs. 6a–c, 7a): praecoxa and intercoxal sclerite of all legs smooth and unarmed. Leg 1 (Fig. 6a): coxa trapezoidal; ornamented with a row of small spinules on antero-distal surface. Basis trapezoidal, somewhat shorter than coxa, ornamented with 1 row of spinules on outer margin and 1 ventral row at base of endopod and another row of small spinules on inner margin, armed with 1 slender seta on outer margin and 1 spiniform seta on inner margin. Exopod 3-segmented, ornamented with 1 row of spinules along outer margins of all segments, second segment with 1 row of spinules at inner distal corner. First segment 0.7 times as long as next 2 segments combined, armed with 1 outer bipinnate spine; second segment unarmed and with 4 elements on third segment (1 outer spine, 1 apical seta and 2 apical geniculate setae). Endopod 2-segmented, about as long as exopod; first segment about as long as first 2 exopodal segments combined, 3.3 times as long as wide, unarmed, ornamented with 1 transverse row of elongate spinules on inner margin and 2 arched rows (1 dorsal, 1 ventral) of large spinules on outer margin; second segment ornamented with 1 row of spinules on outer margin, and armed apically with 1 long geniculate seta and 1 spine; endopodal geniculate seta 1.6 times as long as entire endopod, about as long as innermost geniculate seta on exopod (but almost straight). All exopodal and endopodal armature elements unipinnate along outer margin except bipinnate spine on first exopodal segment. Leg 2 (Fig. 6b): coxa ornamented with 2 arched rows of spinules near outer distal corner (1 dorsal and 1 ventral). Basis rhomboidal, slightly smaller than coxa, unarmed, ornamented with 1 row of large spinules along outer margin. Exopod 3-segmented; ornamented with 1 row of spinules along outer margins of all segments, hyaline frill each at inner distal corner of first and third exopodal segments, but second segment with 1 row of spinules instead. First segment 0.7 times as long as next 2 segments combined, armed with large, bipinnate outer NEW INDIAN SPECIES OF PARASTENOCARIDIDAE

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spine on first segment; second segment unarmed; third segment 1.3 times as long as second segment, armed with 3 long elements, 1 subapical spine and 2 apical bipinnate setae; innermost seta about 1.2 times as long as exopod. Endopod 1-segmented, spatulate and almost 4.1 times as long as wide, 0.6 times as long as first exopodal segment in length, ornamented with 3 large spinules apically, and armed with 1 smooth seta apically, which is 0.7 times as long as segment. Leg 3 (Fig. 7a): coxa rhomboidal, smaller than basis, ornamented with arched row of spinules near inner margin ventrally. Basis robust, ornamented with 1 oblique row of large spinules at distal outer corner, 1 ventral row near inner margin and 1 pore on anterior surface, and armed with moderately long, slender seta on outer margin. Endopod represented by 1 small, curved seta, inserted on inner margin at distal third of basis length. Exopod 2segmented, perfectly fused with each other; ancestral proximal segment moderately strong, nearly thrice as long as basal width and somewhat bent inwards; proximal region somewhat dilated and ornamented with 1 spinule on outer margin and 1 row of 3 spinules at outer distal corner; ancestral second segment (apophysis) short, unornamented, bent inwardly and armed apically with 2 modified, unequal elements (Fig. 7c): inner element ladleshaped with hyaline structure on inner margin, and outer one hyaline, bulbous at base and gradually tapering distally. Thumb distinct at base and modified into curved digitiform chitinized structure, as long as apophysis. Leg 4 (Fig. 6c, d): coxa rhomboidal, ornamented with 1 row of spinules near distal margin. Basis trapezoidal, ornamented with 1 row of spinules and 1 pore on anterior surface, and armed with moderately long seta on outer margin. Exopod 3-segmented, somewhat bent inwards, ornamented with 1 row of long spinules along outer margins of all segments; hyaline frills at inner distal corner of first and third segments, but second segment with 1 row of spinules. First segment nearly 0.7 times as long as next 2 segments combined, armed with single, strong bipinnate outer spine subdistally; second segment 0.7 times as long as third segment; third segment armed with outer spine and long apical, bipinnate seta; seta 1.6 times as long as spine, 2.4 times as long as third exopodal segment, 0.9 times as long as entire exopod. Basal chitinous complex consisting of large sclerotized plate with 1 small hyaline lobe mid-distally and 1 large hyaline lobe at outer distal angle; 1 sturdy, hook-like spine at inner distal corner. Endopod 1.3 times as long as first exopodal segment, membranous and ventricose in outline, with proximal part bulbous, ornamented with 1 row of spinules and drawn out distally into biserrulate, pointed structure. Leg 5 (Figs. 3a, 7d): small, simple, trapezoidal plate, and fused at base; ornamented with 1 group of 2–3 tiny spinules on inner margin, 1 small cuticular pore subproximally; distal part expanded and palmate; inner distal corner produced into rather short spiniform process. Basal seta long, articulate at base and arising from lobed projection; inner lobe with 2 unequal, smooth setae (probably ancestral endopodal armature), outer seta 0.7 times as long as inner seta. Leg 6 (Figs. 2a, 3a): smooth, unarmed, forming simple operculum covering gonopore, fused with sixth pedigerous somite, elliptical in ventral view. Description of adult female. Body length excluding caudal setae 385–405 μm (397 μm in allotype). Habitus (Fig. 4a): ornamentation of prosomites, colour and naupliar eye similar to male, except genital and first abdominal somites fused into double-genital somite. Genital double-somite (Fig. 4b): genital apertures covered by vestigial sixth legs; median copulatory pores also covered by fused sixth legs; seminal receptacles small; copulatory duct very short and weakly sclerotized. Third urosomite, preanal somite, and anal somite very similar to male. Caudal rami (Figs. 3b, 4a, b): 0.7 times as long as anal somite, about 3.2 times as long as wide in dorsal and ventral views, gradually tapering, with armature and ornamentation as in male. Antennule (Figs. 4a, 8e): 7-segmented, first segment short, ornamented with 1 row of spinules on ventral surface; second segment longest; fourth segment with elongate, moderately thick aesthetasc, extending up to the tip of aesthetasc on segment 7; aesthetasc on segment 7 slender and fused basally to 2 apical setae (acrotheck); setal formula: 0.4.2.2+aes.0.0.9+aes. All setae, except unipinnate proximalmost one on segment 2, smooth. Length ratio of antennular segments from proximal to distal end along caudal margin 1.0: 3.0:1.4: 1.7: 0.7: 0.7: 1.5. Antenna, labrum, mandible, maxillule, maxilla and maxilliped similar to male. Leg 1 (Fig. 8a): coxa trapezoidal, with arched row of spinules near proximal outer corner, basis also trapezoidal, with 3 rows of spinules and armed with only outer seta. Other details as illustrated. Leg 2 (Fig. 8b): coxa trapezoidal, ornamented with 1 arched row of spinules at distal inner corner and 1 row of spinules ventrally; basis smaller than coxa, ornamented with 1 row of spinules on outer margin. Exopod same as in male. Endopod somewhat dilated at about midlength; ornamentation and armature same as in male in different views (Fig. 8f–i).

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FIGURE 6. Parastenocaris edakkal n. sp., holotype male: (a) leg 1, anterior view; (b) leg 2, anterior view; (c) leg 4, anterior view; (d) leg 4 endopod, antero-lateral view.

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FIGURE 7. Parastenocaris edakkal n. sp., holotype male (a, d), paratype male (b, c): (a) leg 3, anterior view; (b) leg 6, ventral view; (c) apophysis of leg 3; (d) leg 5, ventral view view.

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FIGURE 8. Parastenocaris edakkal n. sp., allotype female: (a) leg 1, posterior view; (b) leg 2, anterior view; (c) leg 3, posterior view; (d) leg 4, posterior view; (e) antennule, dorsal view; (f–i) leg 2 endopod; (j, k) leg 3 endopod.

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Leg 3 (Fig. 8c): coxa ornamented with 1 arched row of spinules near posterior margin. Basis ornamented with 1 row of spinules on outer margin; armed with 1 long and smooth outer seta. Exopod 2-segmented, ornamented with large spinules along outer margin, both segments with hyaline frill each at inner distal corner; first segment armed with 1 outer spine; second segment with 1 outer spine and 1 apical seta; seta 1.8 times as long as spine; all elements bipinnate. Endopod 1-segmented, slender, almost as long as first exopodal segment, tapering to pointed tip and with spinulose disto-lateral margins. Leg 4 (Fig. 8d): endopod strong, lanceolate, bent inwards, 1.6 times as long as first exopodal segment together with apical biserrulate spine fused at base; 1 large spinule occurring at about mid-inner margin; other details as illustrated. Leg 5 (Fig. 4b, c): fused at base; small, simple, trapezoidal plate, ornamented with 1 small cuticular pore subproximally; subdistal part expanded and inner distal corner produced into moderately long spiniform process. Armature elements same as in male, outer seta 0.8 times as long as inner seta. Leg 6 (Fig. 4b): vestigial, fused into simple cuticular flap, covering gonopores; unornamented and unarmed. Etymology. The specific epithet, alluding to the type locality, Edakkal Cave; proposed here as a noun in apposition to the generic name. Distribution. This species is only known from the type locality. Ecology. Parastenocaris edakkal n. sp. was accompanied by Proserpinicaris corgosinhoi n. sp. and strays of nematodes. This species was collected at a relatively high elevation of 1233 m, whereas its congeneric Indian cavernicole, Parastenocaris kotumsarensis, occurred at an altitude of 560 m. Even Parastenocaris sutlej from a Himalayan river was collected at an altitude of 656 m. Other hyporheic Indian species occur somewhat close to the sea level. Variation. Shape of leg 2 endopod (Fig. 8f–i) varying depending on the angle of view.

Subfamily Fontinalicaridinae Schminke, 2010 Genus Proserpinicaris Jakobi, 1972 sensu Karanovic, Cho & Lee, 2012 Proserpinicaris corgosinhoi n. sp. (Figs 9–14) Type locality. Same as that of Parastenocaris edakkal n. sp. Type material examined. Holotype male, dissected on 2 slides (MNHN-IU-2013-11249), allotype female, dissected on 3 slides (MNHN-IU-2013-11250); 10 paratypes: 1 male (MNHN-IU-2013-11251) and 1 female (MNHN-IU-2013-11252), whole-mounted on 1 slide each, 1 male and 5 females preserved in alcohol (MNHN-IU2013-11253), and 1 male and 1 female in first author’s personal collections; 9 May 2008; Coll. V. R. Totakura & Y. Ranga Reddy. Description of adult male. Total body length, measured from tip of rostrum to posterior margin of caudal rami (excluding caudal setae) 346 μm. Preserved specimens colourless. Naupliar eye absent. Habitus (Fig. 9a) cylindrical and slender; prosome/urosome ratio about 0.8 in dorsal view; greatest width in dorsal view at subdistal cephalothorax. Body length/width ratio about 7.9. Free pedigerous somites without any lateral or dorsal expansions. All segments connected by well-developed arthrodial membranes. Hyaline fringes of all somites smooth, very narrow. Integument sclerotized, smooth, ornamented with sensilla, spinules, pores, and with somewhat subquadrate double-window in posterior half of cephalothorax, and elliptical, slender, dorsal simple cuticular window each on genital and 2 postgenital somites. Pleural areas of cephalothorax and free pedigerous somites well developed; cephalic appendages and coxae of swimming legs clearly exposed in lateral view (not figured). Rostrum (Figs. 9a, 10a) small, linguiform, not demarcated at base, ornamented with 2 large dorsal sensilla, reaching midlength of first antennular segment, about 1.2 times as long as wide. Cephalothorax (Fig. 9a) elongately oval in outline, with subdistal region dilated, about 1.4 times as long as wide in dorsal view and representing 21.5% of total body length. Surface of cephalic shield ornamented with subquadrate integumental double-window and 8 pairs of sensilla. Second and third pedigerous somites as wide as fourth pediger; fourth pediger longer than third one, and free pedigers 2–4 and first urosomite with 2 pairs of posterior sensilla each (Fig. 9a); first urosomite shorter than fourth pediger; genital somite widest of all urosomites, with small elliptical dorsal

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cuticular window in anterior half. Third urosomite as long as second one, with wider elliptical window and 3 pairs of sensilla. Fourth urosomite about as long as second urosomite, with dorsal cuticular window and 3 pairs of large posterior sensilla. Preanal somite narrower and slightly shorter than fourth urosomite, with largest dorsal cuticular window, extending latero-ventrally. Anal somite about as long as wide and slightly narrower than preanal somite, ornamented with 1 pair of large dorsal sensilla near base of anal operculum, 1 pair of large lateral cuticular pores in anterior half and 1 row of spinules on either side proximo-ventrally. A single large, longitudinally placed spermatophore discernible through cuticle of fifth and genital somites, about 2.1 times as long as wide, beanshaped, with narrow neck. Anal operculum well developed, smooth and bowl-shaped, with slightly concave distal margin, representing 59.1% of somite's width. Anal sinus wide and smooth. Caudal rami (Fig. 9a–c): short, about 1.3 times as long as greatest width in dorsal view, 1.2 times in lateral view (Fig. 9b), and about 0.4 times as long as anal somite, proximal third slightly dilated; divergent, with space between them being about 1.2 times as long as that of 1 ramus maximum width; armed with 6 elements (2 lateral, 1 dorsal, 1 subapical, 2 apical); and ornamented with 1 row of spinules, and 1 cuticular pore disto-laterally; setae I and III thin, unequal, inserted at 2/5 of ramus; seta II missing. Dorsal seta (VII) slender and inserted close to inner margin in distal half, about 1.8 times as long as caudal ramus, biarticulate basally. Inner apical seta (VI) smooth, proximally swollen, inserted close to ventral margin, about 1.5 times as long as ramus length. Middle apical seta (V) strongest, without breaking plane, bipinnate, about 8.8 times as long as ramus, pointing distally, with straight, acute tip. Outer subapical seta (IV) without breaking plane and unipinnate, about 2.2 times as long as ramus, inserted close to dorsal surface and pointing laterally. Antennule (Fig. 10b, c): 1.2 times as long as cephalothorax, stout, 8-segmented, strongly prehensile and digeniculate, proximal geniculation between third and fourth, and distal geniculation between sixth and seventh segments. First segment short, ornamented with 1 row of spinules. Second segment relatively short, about as long as fifth segment. Segments 4–5 much dilated, fifth segment with elongate, aesthetasc with narrow, blunt tip, reaching midlength of ultimate segment, fused basally to simple seta; apical aesthetasc on eighth segment slender, shorter than segment, fused basally with 2 setae (acrotheck). Setal formula: 0.6.4.2.8+aes.2.0.9+aes. All setae slender, smooth except proximalmost seta on second segment unipinnate with long setules. Length ratios of antennular segments from proximal to distal end along caudal margin 1:2.3:1.1:0.8:2.3:1.1:1.3:1.1. Antenna (Fig. 10d): composed of coxa, allobasis, 1-segmented endopod, and 1-segmented exopod. Coxa short, ornamented with 1 row of short spinules, unarmed. Allobasis about 3.2 times as long as maximum width, unarmed, ornamented with 2 rows of spinules on inner margin. Exopod small, cylindrical, about thrice as long as wide, unornamented, armed with 1 unipinnate apical seta, which is about twice as long as segment. Endopod 0.7 times as long as allobasis and about twice as long as wide, with surface frill distally, ornamented with 2 longitudinal rows of spinules on inner margin and 2 rows of spinules distally; and armed with 2 unequal, bipinnate spines laterally and with 5 strong elements apically (2 spines, 2 geniculate setae and 1 unipinnate transformed seta). Labrum large and subtriangular in lateral view, other details not studied (not figured). Mandible (Fig. 10e): with narrow cutting edge on elongated coxa, armed with 2 complex teeth ventrally, 1 unipinnate seta dorsally, and several smaller teeth. Palp 1-segmented, cylindrical, about 3.2 times as long as wide, unornamented and armed with 2 smooth, unequal apical setae. Maxillule (Fig. 10f): arthrite sub-rectangular, about 2.2 times as long as wide in lateral view, armed with 1 strong lateral seta and 3 apical elements. Coxal endite armed with 1 smooth apical seta. Basis twice as long as coxal endite, armed with 2 smooth apical setae. Maxilla (Fig. 10g): composed of syncoxa, basis, and 1-segmented endopod. Syncoxa with 2 endites; basal one short, armed with 1 smooth seta apically, distal one long, armed with 1 smooth and 1 pinnate setae apically. Allobasis prolonged into strong unipinnate claw, without seta at base; endopod smallest, armed with 2 setae. Maxilliped (Fig. 10h): with short syncoxa, unarmed and unornamented; basis slender, 2.3 times as long as wide, unornamented and unarmed; endopod small with unipinnate claw. Swimming legs (Figs. 11a–b, 12a–d): praecoxa and intercoxal sclerite of all legs smooth and unarmed.

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FIGURE 9. Proserpinicaris corgosinhoi n. sp., paratype male (a), holotype male (b, c): (a) habitus, dorsal view; (b) urosome, lateral view; (c) urosome, ventral view.

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FIGURE 10. Proserpinicaris corgosinhoi n. sp., holotype male (a–h); allotype female (i): (a) rostrum, latero-ventral view; (b) antennule, ventral view; (c) antennule, lateral view (ornamentation and armature omitted); (d) antenna, lateral view; (e) mandible, lateral view; (f) maxillule, lateral view; (g) maxilla, lateral view; (h) maxilliped, lateral view; (i) antennule, lateral view.

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FIGURE 11. Proserpinicaris corgosinhoi n. sp., holotype male (a–b), paratype male (c), paratype female (d): (a) leg 1, anterior view; (b) leg 2, posterior view; (c) leg 5, ventral view; (d) leg 5, ventral view.

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FIGURE 12. Proserpinicaris corgosinhoi n. sp., holotype male (a, d), paratype male (b, c, e, f): (a) leg 3, anterior view; (b) leg 3, apophysis, anterior view; (c) leg 3, thumb, anterior view; (d) leg 4, posterior view; (e, f) leg 4 endopod, posterior view.

Leg 1 (Fig. 11a): coxa unarmed, ornamentation not discernible. Basis trapezoidal, ornamented with 1 row of large spinules on outer margin, and 1 row at base of endopod ventrally, armed with small, smooth outer seta. Exopod 3-segmented, ornamented with 1 row of spinules each along outer margins of all segments; armed with outer bipinnate spine on first segment. First segment 0.6 times as long as next 2 segments combined; second segment unarmed; 4 elements on third segment (1 outer spine, 1 apical seta and 2 apical geniculate setae). Endopod 2-segmented, first segment 3.3 times as long as wide, unarmed, ornamented with 1 row of spinules on inner margin and 1 ventral row of large spinules on outer margin, 1 row distally; second segment ornamented with spinular row on distal margin, armed apically with 1 outer spine and 1 long geniculate seta; endopodal geniculate seta 1.4 times as long as entire endopod, almost twice as long as outer spine, about as long as inner geniculate seta on exopod. All exopodal and endopodal armature elements unipinnate along outer margins except bipinnate spine on first exopodal segment. NEW INDIAN SPECIES OF PARASTENOCARIDIDAE

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Leg 2 (Fig. 11b): coxa ornamented with 1 arched row of spinules near outer margin and 1 oblique row medially near distal margin. Basis slightly smaller than coxa, unarmed, ornamented with 1 row of large arched spinules at distal outer corner and 1 pore proximally. Exopod 3-segmented; ornamented with 1 row of spinules each along outer margins of all segments, hyaline frill each at inner distal corner of first and third exopodal segments but second segment with 1 row of spinules instead. First segment 0.6 times as long as next 2 segments combined, armed with large, outer spine on first segment; second segment unarmed; third segment slightly longer than second segment, armed with 3 long elements (1 subapical unipinnate spine and 2 apical bipinnate setae); inner apical seta about 1.2 times as long as exopod. Endopod 1-segmented, cylindrical, and 3.2 times as long as wide, 0.4 times as long as first exopodal segment, ornamented with 2 large spinules apically on distal margin, 3 small spinules on inner margin and 2 spinules at midlength ventrally; armed apically with 1 smooth, straight seta, which is 0.9 times as long as segment. Leg 3 (Fig. 12a–c): coxa trapezoidal, ornamented with arched row of ventro-medial spinules close to middistal margin. Basis produced disto-medially into 1 lobed structure; ornamented with 1 oblique row of long spinules and 1 pore on anterior surface. Endopod represented by a short spiniform seta at about proximal third of basis length. Exopod modified, both segments fused; ancestral proximal segment stumpy, with almost straight outer margin, 1.5 times as long as wide, about as long as basis, somewhat bent inwards, ornamented with 1 row of 3 spinules on distal outer margin. A massive tubular pore arising from proximal anterior surface. Ancestral second segment mace-like in anterior view (Fig. 12b), membranous, dilated subdistally, with irregular lateral margins, unornamented. Thumb dagger-shaped, gradually tapering, with acuminate tip and irregular hyaline membrane laterally (Fig. 12c). Leg 4 (Fig. 12d–f): coxa trapezoidal, ornamented with 1 row of spinules near distal margin. Basis longer than coxa and trapezoidal in posterior view, ornamented with 1 row of spinules on outer margin and 1 pore proximally, armed with moderately long seta on outer margin. Exopod relatively short and thick, 3-segmented; ornamented with 1 row of spinules each along outer and inner margins of all segments; inner spinules on first segment particularly long; hyaline frill at inner distal corner of third segment. First segment 0.7 as long as next 2 segments combined, armed with 1 strong bipinnate outer spine subdistally; second segment unarmed and with straight inner margin; third segment slightly longer than second one, armed with 1 subapical spine and 1 apical bipinnate seta; apical seta 1.8 times as long as outer spine, about 1.7 times as long as third exopodal segment, half as long as entire exopod. Endopod short, membranous and somewhat conical, and additional hyaline structure relatively large, foliaceous and occurring rather close to, and partially overlapping, endopod. Leg 5 (Figs. 9b, c, 11c): both legs fused at base, with narrow space proximally; represented by subquadrate plate, with distal inner corner rounded; ornamented with 1 row of spinules along distal half of inner margin, gradually increasing in size distally and 1 small cuticular pore on proximal surface; armed with 3 elements: 1 long articulate basal seta and 1 outer smooth, outcurved, simple seta, slightly shorter than inner spiniform seta (possible ancestral endopod). Leg 6 (Fig. 9b, c): smooth, unornamented, forming simple operculum covering gonopore, fused with sixth pediger, and plate-like structure in ventral view. Description of adult female. Body length, excluding caudal setae 384 μm. Habitus (Fig. 13a): ornamentation of prosomites, colour and naupliar eye similar to male, except genital and first abdominal somites fused into double-genital somite. Genital double-somite: without any trace of subdivision, with oval dorsal cuticular window (Fig. 13b) in anterior half, ornamented with 1 sensillum on either side of cuticular window (Fig. 13b). Genital complex occupying anterior ventral half of genital double-somite; genital apertures paired, covered by vestigial sixth legs; copulatory pores median; seminal receptacles small, hard to distinguish from internal tissue and gut content; copulatory duct very short and weakly sclerotized. Preanal somite, anal somite and urosomites very similar to male. Caudal rami (Fig. 13a–c): divergent, 0.4 times as long as anal somite, about 1.2 times as long as wide in lateral view (Fig. 13b), 1.4 times in dorsal and ventral views (Fig. 13a, c), with armature and ornamentation as in male, but inner apical seta (VI) slightly swollen at base.

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FIGURE 13. Proserpinicaris corgosinhoi n. sp., paratype female, (a, b); allotype female (c): (a) habitus, dorsal view; (b) urosome, lateral view; (c) urosome, ventral view.

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FIGURE 14. Proserpinicaris corgosinhoi n. sp., allotype female: (a) leg 1, posterior view; (b) leg 2, posterior view; (c) leg 3, posterior view (arrowhead pointing to spinular row); (d) leg 4, posterior view.

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FIGURE. 15. Proserpinicaris karanovici n. sp., (a) paratype male, habitus, lateral view; (b) holotype male, urosome, ventral view.

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FIGURE 16 Proserpinicaris karanovici n. sp., holotype male (a–i); allotype, female (j): (a) rostrum, dorsal view; (b) antennule, ventral view; (c) antenna, lateral view; (d) labrum, ventral view; (e–f) mandible, lateral view; (g) maxillule, lateral view; (h) maxilla, lateral view; (i) maxilliped, lateral view; (j) antennule, lateral view.

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FIGURE 17. Proserpinicaris karanovici n. sp., holotype male: (a) leg 1, anterior view; (b) leg 2, posterior view; (c) leg 3, anterior view; (d) leg 4, anterior view; (e) hyaline structure on basis of leg 4; (f) leg 5, ventral view.

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Antennule (Fig. 10i): 7-segmented, first segment with few minute spinules at outer distal corner in lateroventral view; slender, elongate aesthetasc on fourth segment reaching almost distal margin of ultimate segment; seventh segment with relatively slender and short apical aesthetasc, which is fused basally to 2 apical setae; setal formula: 0.4.4.3+aes.0.0.8+aes. All setae smooth except unipinnate proximalmost one on second segment. Proximal seta on second and ultimate segment with basal articulation. Length ratio of antennular segments, from proximal to distal end and along caudal margin 1:2.7:1.6:1.8:1.1:1.1:1.6. Antenna, labrum, mandible, maxillule, maxilla, maxilliped, and leg 1 similar to male. Leg 2 (Fig. 14b): exopodal segments same as in male. Endopod cylindrical, armed with seta, which is 1.5 times as long as segment and reaching distal margin of first exopodal segment; ornamented with 1 spinule on outer margin, and 3 spinules on distal margin. Leg 3 (Fig. 14c): coxa with arched spinular row on anterior surface. Basis ornamented with 1 row of spinules on outer margin and 1 row near base of endopod and 1 pore; armed with very long and smooth outer seta, articulate at base, 1.2 times as long as entire exopod. Exopod 2-segmented, ornamented with large spinules along outer margin and 1 row at inner distal corner of first segment; second segment alone with hyaline frill at inner distal corner; first segment armed with outer spine; second segment with 1 outer spine and 1 apical strong seta; seta 1.6 times as long as spine; all elements bipinnate. Endopod small, 2.6 times as long as wide, only half as long as first exopodal segment tapering to pointed tip, ornamented with transverse row of tiny spinules subdistally. Leg 4 (Fig. 14d): exopod similar to male. Endopod rather slender, lanceolate, 1.5 times as long as first endopodal segment, with apical spine fused at base and having sparse spinules on lateral margins; 1 circlet of 3 spinules occurring at midlength (base of fused apical spine), and 1 spinule on proximal outer margin. Leg 5 (Figs. 11d, 13b, c): both legs fused at base with narrow inter-space proximally; each in the form of obovate plate, reaching midlength of next segment and ornamented with 1 row of spinules on inner margin, spinules increasing in size distally; basal seta long, articulate at base; distal margin armed with 1 bare outer seta and 1 bipinnate inner spine, seta about as long as spine; distal inner corner produced into somewhat elongate inner spiniform process. Sixth leg vestigial. Etymology. The new species is named in honour of Dr. P. H. C. Corgosinho, Universitário Frutal-MG in Brazil, a well-known authority on the Neotropical Parastenocarididae. The specific epithet is a noun in the genitive singular. Gender masculine. Distribution. This species is only known from the type locality. Ecology. This species was accompanied by Parastenocaris edakkal n. sp. and nematodes in very small numbers.

Proserpinicaris karanovici n. sp. (Figs. 15–20) Type locality. Farm bore in the riparian zone of the River Krishna at Kunchanapalli village (16º23′42.1″N, 80º32′28.2″E, elevation 8.9 m; water temperature 26ºC, pH 7.0), 3 km from Vijayawada city in Guntur District, Andhra Pradesh (Fig 1a). Type material examined. Holotype male (MNHN-IU-2013-11935) and allotype female (MNHN-IU-201311936), dissected on 3 slides each; 4 paratypes: 1 paratype male (MNHN-IU-2013-11937), dissected on 2 slides; 1 male (MNHN-IU-2013-11938) and 2 females (MNHN-IU-2013-11939–11940), whole-mounted on 1 slide each. 4 January 2010, Coll. V. R. Totakura. Other material examined. India, Andhra Pradesh, Guntur District, 1 male and 2 females collected from the type locality; 15 December 2013, Coll. V. R. Totakura. Description of adult male. Total body length, measured from tip of rostrum to posterior margin of caudal rami (excluding caudal setae), 522 μm. Preserved specimens colourless. Naupliar eye absent. Habitus (Fig. 15a) with thin cuticle, smooth and not pitted, cylindrical and slender, without any podoplean demarcation between prosome and urosome; prosome/urosome ratio about 0.8 in lateral view; greatest width at fifth pedigerous somite in lateral view; free pedigerous somites 2–4 gradually increasing in size. Body length/width ratio about 8.3. Cephalothorax about as wide as genital somite in lateral view, 18.9% of body length. Free pedigerous somites with narrow

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arthrodial membranes. Hyaline fringes of all somites smooth, very narrow and hard to distinguish from arthrodial membranes. Integument smooth, ornamented only with sensilla, spinules and with single rectangular dorsal cuticular window on cephalothorax and vague elliptical cuticular window each on urosomites 2–5. Pleural areas of cephalothorax and free pedigerous somites present, cephalic appendages and coxae of swimming legs partly exposed in lateral view (Fig. 15a). Rostrum (Fig. 16a) small, membranous, about as long as wide, linguiform, fused at base, ornamented with 2 large dorsal sensilla, reaching about end of first antennular segment. Cephalothorax (Fig. 15a) subquadrate in lateral view, about 1.5 times as long as wide in lateral view and 21.3% of total body length. Surface of cephalic shield ornamented with sensilla and 1 elliptical, smooth window in distal half; 4 pairs of sensilla around window; 3 pairs of sensilla on either side and 1 pair at base of antennule. Second pedigerous somite 0.9 times as wide as posterior half of cephalothorax in dorsal view, with 3 pairs of large sensilla (1 dorsal and 1 lateral, 1 ventral). Third pedigerous somite slightly wider than second one, with 3 pairs of large sensilla. Fourth pedigerous somite slightly wider and longer than third prosomite, with only 3 pairs of large posterior sensilla. Urosome (Fig. 15a, b): urosomites gradually narrowing posteriorly, with poorly developed dorsal hyaline frills, hard to distinguish from arthrodial membranes between urosomites. Urosomites 2–5 each with elliptical cuticular window, and 1–3 each ornamented with 4 pairs of large posterior sensilla (2 dorsal, 1 lateral, 1 ventral). Fourth urosomite with 3 pairs of large posterior sensilla. Preanal somite slightly narrower and longer than fourth urosomite, without any surface ornamentation in dorsal view. Anal somite about as long as wide and slightly narrower than preanal somite, ornamented with 2 large dorsal sensilla at base of anal operculum, 1 cuticular pore on either side and 1 row of spinules on ventro-distal margin, of which 1 spinule large. A large, longitudinally placed spermatophore (Fig. 15a) visible through cuticle of fifth pediger and genital somite, about 3.1 times as long as wide, bean-shaped, with narrow and curved neck. Anal operculum well developed, distal margin convex, not reaching posterior end of anal somite, representing 68% of somite's width. Anal sinus wide; ornamented with 2 diagonal rows of slender spinules. Caudal rami (Fig. 15a, b): subcylindrical, sub-proximally dilated on inner margin and gradually tapering, about 2.6 times as long as greatest width in ventral view, 2.2 times in lateral view, and about 1.2 times as long as anal somite, slightly divergent, with space between them being about half the maximum width of ramus; armed with 6 setae (seta II absent) and ornamented with posterior row of spinules on ventral margin and 1 disto-lateral cuticular pore. Lateral setae (I and III) located at 2/5 of ramus length, unequal. Dorsal seta (VII) inserted at about 3/4 of, and close to, inner margin, nearly 1.3 times as long as caudal ramus, plumose, biarticulate basally. Inner apical seta (VI) smooth, inserted close to ventral margin, about 0.7 times as long as ramus. Middle apical seta (V) strongest, without breaking plane, bipinnate, about 2.4 times as long as ramus, pointing distally, with straight tip. Outer apical seta (IV) greatly reduced, smooth, only about 0.3 times as long as ramus length, inserted close to dorsal surface and pointing postero-laterally. Antennule (Fig. 16b): 8-segmented, slightly longer than cephalothorax, slender, prehensile, strongly digeniculate, geniculation between third and fourth, and sixth and seventh segments; last 2 segments directed distally (‘coiled type’). First segment very short, ornamented with 1 row of tiny spinules. Segments 3–5 moderately dilated, fifth segment without any proximal spiniform process on anterior surface; aesthetasc elongate with blunt tip, reaching mid-length of ultimate segment, fused basally to simple seta; short and slender apical aesthetasc on ultimate segment, fused basally to 2 setae (acrotheck). Setal formula: 0.6.4.1.5+aes.2.0.9+aes. All setae slender and smooth except 1 spiniform seta on third segment; proximalmost seta on second segment unipinnate, with extremely long setules along outer margin. Length ratios of antennular segments, along medial axis, 1.0: 3.1: 1.1: 0.3: 1.2: 1.1: 1:1. Antenna (Fig. 16c): composed of coxa, allobasis, 1-segmented exopod and 1-segmented endopod. Coxa very short, unarmed and unornamented. Allobasis about 3.5 times as long as maximum width, unarmed but ornamented with 2 arched rows of spinules on anterior surface. Exopod slender, cylindrical, about 5 times as long as wide, unornamented, and armed with 1 unipinnate apical seta, which is 2.3 times as long as segment. Endopod 0.4 times as long as allobasis and about twice as long as wide, with surface frill subdistally, ornamented with 2 longitudinal spinular rows on inner margin, armed laterally with 2 short, bipinnate, unequal spines and apically with 5 strong elements (2 spines, 2 geniculate setae and 1 unipinnate transformed seta). Labrum (Fig. 16d): large and subtriangular in ventral view, with narrow and arched cutting edge, and 1 row of elongate spinules on ventral surface.

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Mandible (Fig. 16e, f): gnathobase with narrow cutting edge on elongated coxa, armed with 2 complex teeth ventrally, 1 unipinnate seta dorsally, and several smaller teeth. Palp 1-segmented, cylindrical, about 4.2 times as long as wide, unornamented, armed apically with 2 smooth apical setae. Maxillule (Fig. 16g): with relatively large praecoxa; praecoxal arthrite about 3.2 times as long as wide; no ornamentation on posterior surface; armed with 3 strong, smooth, apical spinous processes and 1 subapical stout spiniform seta. Coxal endite armed with 2 smooth setae apically. Basis slightly longer than coxal endite, armed with 3 smooth apical setae. Maxilla (Fig. 16h): composed of syncoxa, basis, and 1-segmented endopod. Syncoxa with 2 endites, basal one short, armed with 1 apical smooth, fused seta; distal endite armed with 1 smooth seta and 1 unipinnate seta apically. Allobasis prolonged into strong unipinnate claw, without seta at base. Endopod short, with 2 unequal setae. Maxilliped (Fig. 16i): syncoxa short and relatively strong, unarmed and unornamented; basis slender, 5.4 times as long as wide, unornamented and unarmed; endopod small, with unipinnate claw, about 0.5 times as long as basis. Leg 1 (Fig. 17a): coxa with 1 circlet of spinules on posterior surface. Basis smaller than coxa, ornamented with 1 row of large spinules along outer margin and another row of spinules between exopod and endopod ventrally, and 1 pore proximally; armed with 1 slender, short seta on outer margin. Exopod 3-segmented; first segment armed with 1 outer bipinnate spine, 0.9 times as long as next 2 segments combined and 4 elements on third segment (1 outer spine, 1 apical seta and 2 apical geniculate setae); ornamented with few large spinules along outer margin. Endopod 2-segmented, longer than exopod; first segment longer than proximal 2 exopodal segments combined, ornamented with 1 row of elongate spinules on subdistal inner margin, 2 rows of short spinules on outer margin; second endopodal segment ornamented with 1 row of spinules on mid-outer margin and armed apically with 1 long geniculate seta and 1 short spine; geniculate seta 1.2 times as long as entire endopod, almost 2.1 times as long as outer spine on endopod, about as long as inner geniculate seta on exopod. All exopodal and endopodal armature elements unipinnate along outer margin except bipinnate spine on first exopodal segment. Leg 2 (Fig. 17b): coxa large, ornamented with 1 transverse row of small spinules on posterior outer margin. Basis ornamented with 1 row of spinules near anterior inner margin, 1 pore on mid-dorsal surface and unarmed. Exopod 3-segmented; ornamented with 1 row of spinules along outer margins of all segments and hyaline frill at inner distal corner of first and third segments; second segment with 1 row of spinules at inner distal corner; first segment 0.8 times as long as next 2 segments combined and armed with 1 outer bipinnate spine; second segment unarmed; third segment as long as second one, armed with 3 long setae (1 subapical unipinnate and 2 apical bipinnate setae); innermost seta about as long as exopod. Endopod 1-segmented, cylindrical and almost 6 times as long as wide, reaching 1/3 of first exopodal segment in length, ornamented with 2 spinules on distal end, and armed apically with 1 smooth seta, which is about as long as segment and pointing outwards. Leg 3 (Fig. 17c): characteristically straight; coxa trapezoidal, ornamentation not discernible. Basis rectangular but produced into 1 lobe-like structure at inner distal corner in anterior view; ornamented with 1 oblique row of unequal spinules near outer margin, 2 rows of small spinules near inner proximal margin, 1 pore proximally; armed with long, slender outer seta, which is articulate at base. Endopod represented by short, sturdy spiniform structure, inserted at 2/3 of inner margin. Exopod modified, fused; proximal ancestral segment subquadrate; ornamented with 1 row of 3 small spinules on outer distal margin; apophysis doubly curved, claw-like and unornamented; thumb dagger-shaped, membranous, distinct at base, slightly dilated subproximally, and 1.5 times as long as apophysis; 1 narrow hyaline membrane at about mid-inner margin on lateral margins. Leg 4 (Fig. 17d, e): coxa rhomboidal, ornamented with 1 row of spinules posteriorly. Basis trapezoidal, ornamented with 1 row of spinules on anterior surface, 1 row on outer proximal margin, and 1 pore on proximal surface; armed with moderately long outer seta. Exopod slender, elongate, 3-segmented, ornamented with spinules only along outer margin; first segment 0.6 as long as next 2 segments combined; hyaline frill at inner distal corner of first and third segments, second segment with 1 row of spinules at inner distal corner; first segment armed with strong bipinnate outer spine subdistally; second segment unarmed; third segment slightly longer than second one and armed with 1 outer spine and 1 inner, apical bipinnate seta; seta 2.7 times as long as spine, 2.3 times as long as third exopodal segment, 0.8 times as long as entire exopod. Endopod nearly as long as first exopodal segment, membranous, proximal 2/3 slightly swollen; lateral margins fringed with tiny spinules; the hyaline structure leaflike and occurring close to, and partially overlapping, endopod . Leg 5 (Figs. 15a, 17f): both legs separate at base; each leg large, elongate trapezoidal plate, ornamented with longitudinal row of 7 large spinules on distal inner margin, spinules increasing in size posteriorly, distalmost

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spinule slightly overreaching tip of inner spiniform process; and 1 small cuticular pore on proximal half; inner distal angle protruding as spiniform process and with minute spinules distally, spiniform processes of both legs divergent; each leg armed with 4 smooth setae on uneven outer margin. Basal lobe distinct, carrying 1 long, basally articulate seta followed by 1 tiny seta; next 2 setae unequal and arising from a slight bulge (probably ancestral endopod). Leg 6 (Fig. 15a): claw-like, bare, posteriorly directed, forming simple operculum covering gonopore, fused with sixth pedigerous somite. Description of adult female. Body length excluding caudal setae 540 μm. Habitus (Fig. 18a): ornamentation of prosomites and colour similar to male, except genital somite and first abdominal somite fused into doublesomite. Cephalothorax (Fig. 18a) elongate-oval with postero-lateral corners dilated in dorsal view, about 1.2 times as long as wide in dorsal view; representing 15.9% of total body length. Genital double-somite about as long as wide without any trace of subdivision and with vague, elliptical cuticular window on antero-dorsal half, and 4 pairs of large posterior sensilla. Other urosomites gradually decreasing in size. Preanal and anal somites same as in male. Genital complex occupying antero-ventral half of genital double-somite; genital apertures paired, each covered by vestigial sixth legs; copulatory pores medial; seminal receptacles small; copulatory duct very short and weakly sclerotized. Caudal rami (Figs. 18a, 20a, b): 1.3 times as long as anal somite, about 2.8 times as long as wide; gradually tapering; armature and ornamentation as in male. Antennule (Fig. 16j): 7-segmented, first segment with few minute spinules near outer margin, fourth segment with slender aesthetasc, overreaching midlength of ultimate segment, and more slender apical aesthetasc on seventh segment, which is fused basally to 2 apical setae; setal formula: 0.4.5.2+aes.1.1.8+aes. All setae, except proximalmost one on second segment smooth. Length ratios of antennular segments on medial axis 1.0: 3.5: 1.5: 1.3: 0.7: 1.0: 1.2. Antenna, labrum, mandible, maxillule, maxilla, maxilliped and leg 1 similar to male. Leg 2 (Fig. 19b): exopod same as in male. Endopod 1-segmented, slender, 6.3 times as long as wide, somewhat curved, ornamented with 1 spinule subdistally; apex with 2 spinules and 1 slender seta, reaching distal end of first exopodal segment. Leg 3 (Fig. 19c): coxa with arched row of spinules on anterior surface. Basis ornamented with spinules on outer margin and 1 pore anteriorly, armed with long and smooth outer seta, which is about as long as entire exopod. Exopod 2-segmented, ornamented with 1 row of large spinules along outer margin, and both segments with hyaline frill each at inner distal corner. First segment armed with 1 outer spine; second one with 1 outer spine and 1 apical strong seta; seta 2.9 times as long as spine; all elements bipinnate. Endopod 1-segmented, cylindrical, blunt, 4.4 times as long as wide, short, 0.3 times as long as first exopodal segment, ornamented with 1 spinule at midlength on inner margin and 1 spinule apically, which is 0.3 times as long as segment. Leg 4 (Fig. 19d): exopod similar to male. Endopod lanceolate, about as long as first exopodal segment together with fused apical spine, somewhat curved inwards, ornamented with 2 circlets of spinules in proximal half. Leg 5 (Figs. 19e, 20a, b): trapezoidal plate, overreaching posterior margin of fifth pediger; inner margin ornamented with only 3 small spinules, with distinct gap between proximal spinule and distal pair of spinules. Inner spiniform processes on both legs parallel; setal armature same as in male. Leg 6 completely absent. Etymology. The new species is named in honour of Prof. T. Karanovic, Hanyang University in Seoul, in recognition of his significant contributions to the systematics of groundwater copepods. The specific epithet, karanovici, is a noun in the genitive singular. Distribution. The new species is confined to the type locality only. Ecology. The new species was accompanied by various other taxa on two occasions at the type locality: Atopobathynella sp., Habrobathynella sp., Serbanibathynella sp., an unidentified cyclopoid, Nitocrella sp., Parastenocaris sp., an unidentified harpacticoid, water mites, nematodes, oligochaetes and insect larvae on 4 January 2010; and Atopobathynella sp., Serbanibathynella sp., Nitocrella sp., Parastenocaris sp., and an unidentified harpacticoid on 15 December 2013. Similar multiple stygobitic species coexistence has been reported elsewhere (see Corgosinho & Martínez Arbizu 2005; Corgosinho et al. 2007; Karanovic & Cooper 2011). Variation. Caudal rami varying in size and form in female (Fig. 18a–e).

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FIGURE 18. Proserpinicaris karanovici n. sp., paratype female: (a) habitus, dorsal view; (b–d) caudal ramus, lateral view; (e) anal somite and caudal rami, latero-ventral view.

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FIGURE 19. Proserpinicaris karanovici n. sp., allotype female: (a) leg 1, posterior view; (b) leg 2, anterior view; (c) leg 3, anterior view; (d) leg 4, anterior view; (e) leg 5, ventral view.

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FIGURE 20. Proserpinicaris karanovici n. sp., allotype female: (a) urosome, ventral view; (b) urosome, lateral view.

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Discussion That Parastenocaris edakkal n. sp. belongs to the brevipes-group, as proposed by Lang (1948) and revised by Reid (1995), is very clear, inter alia, from the characteristic endopodal complex of the male leg 4; the long, distally serrulate female leg 4 endopod, and the tapering caudal rami of both sexes with dorsal and lateral setae inserted at about midlength (see Karanovic 2005). Karanovic (2005) discussed this group and provided a key for 14 valid species. Following the later addition of one species each from Australia and South Korea (Parastenocaris jane Karanovic, 2006 and Parastenocaris koreana Karanovic & Lee, 2012), the revalidation of the Japanese Parastenocaris biwae Miura, 1969, and the allocation of two originally ungrouped species (viz. Parastenocaris palmerae Reid, 1991, from USA and Parastenocaris muvattupuzha Ranga Reddy & Defaye, 2009, from India), the current tally of recognised species in the brevipes-group has increased to 18 (see Karanovic & Lee 2012a). Karanovic & Lee (2012a) provided a key for the males of all these species and further revised it in their monograph on Parastenocarididae (see Karanovic & Lee 2012b). Parastenocaris edakkal n. sp. has closest affinity with the Sri Lankan Parastenocaris brincki rather than with any of the known Indian congeners, despite the fact that the morphology of P. brincki is incomplete and its female still unknown. Parastenocaris edakkal n. sp. shares with P. brincki the following salient features: leg 4 endopod proximally bulbous and distally tapering off to acute point, and bearing one or more spinules, and also ornamented with one diagonal row spinules at about midlength; basis of the same appendage ornamented with one arched row of spinules; leg 5 with short the spiniform process at the inner distal corner albeit slightly longer in the latter; and leg 3 apophysis bearing two apical elements. However, the two species are distinctly different from each other in several other respects: anal somite 1.48 is times as long as vs. same as caudal ramus; leg 3 exopod slender and long vs. short and stout; without vs. with triangular hyaline membrane on outer distal margin; and the thumb is ladle-like vs. hook-like; leg 4 basis bearing strong, claw-like vs. slender, nearly straight inner tooth; the chitinous complex has two hooks vs. one hook; leg 5 inner margin is ornamented with two or three spinules vs. unornamented, and armed with three vs. two setae on outer margin; and caudal ramus with three vs. two lateral setae. Parastenocaris edakkal n. sp. is distinctly different from all other members of the brevipes-group by a combination of the following characters: body length of males 368–382 μm and of females 386–405 μm; body heavily chitinised and pitted; male urosomites 2–5 and female urosomites 2–4 with dorsal, elliptical integumental window each; caudal rami of both sexes about 3.2 times as long as wide, somewhat tapering posteriorly; lateral setae (I–III) inserted slightly distal to midlength of ramus; outer apical seta (IV) about as long as ramus; in male, aesthetasc on antennular segment 5 much elongate, overreaching ultimate segment; leg 1 basis with inner seta; leg 3 apophysis about as long as thumb with two modified apical elements; leg 4 basal chitinous complex consisting of large sclerotized plate with two accessory hooks or lobes on posterior border and one sturdy hook-like spine at inner distal corner; endopod membranous, proximal part bulbous and ornamented with one oblique row of four or five spinules, and drawn out distally into biserrulate, pointed structure; leg 5 sexually dimorphic; male leg 5 palmate, with short spiniform process at inner distal corner and a group of two or three spinules on inner margin; female leg 5 trapezoidal, with distal inner corner produced into smooth, relatively long spiniform process; both legs armed with three setae on distal margin; and female leg 4 endopod spiniform, reaching distal margin of second exopodal segment. In this context, it seems appropriate to reexamine the taxonomic relationships of the as-yet ungrouped Indian Parastenocaris kotumsarensis Ranga Reddy & Defaye, 2009, which is apparently a ‘puzzling’ species within the genus Parastenocaris (see Ranga Reddy & Defaye 2009), and to update the world list of the brevipes-group of Parastenocaris species. Parastenocaris kotumsarensis was listed under incertae sedis of the subfamily Parastenocaridinae by Schminke (2010). Though Ranga Reddy & Defaye (2009) did point out the close affinity of this species with the brevipes-group based on the male leg 4 endopodal complex, which no doubt is the signal synapomorphy of this group, they refrained from assigning it to this group because of its sexually isomorphic fifth legs together with the presence of ventro-lateral integumental windows on the penultimate and antepenultimate urosomites, and the relatively short female leg 4 endopod. Now that the lack of sexual dimorphism in the fifth legs could be considered a plesiomorphic character in a larger group of species (Schminke 2010; Karanovic & Lee 2012b), it would perhaps be appropriate to regard the other two peculiar features as autapomorphies of the species, thus paving the way for its allocation to the brevipes-group. Two other Indian species, viz. Parastenocaris sutlej Ranga Reddy, 2011 and Parastenocaris gundlakamma Ranga Reddy, 2011 (see Ranga Reddy 2011a) rightly

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belong to the brevipes- group. Following the inclusion of the afore-said Indian species in the brevipes-group, the total number of world species of this group goes up to 21. Most parastenocaridids lack an inner spine/seta on the leg 1 basis (Galassi & De Laurentiis 2004), but its presence in the males has up till now been observed not only in P. edakkal n. sp. but in four other Indian species as well, viz., P. savita, P. muvattupuzha, P. gundlakamma and P. tirupatiensis Ranga Reddy, 2012 (see Ranga Reddy & Defaye 2009; Ranga Reddy 2001, 2011a, 2012). According to Galassi & De Laurentiis (2004), the inner seta on the leg 1 basis of females and/or males could be present in certain species belonging to different phylogenetic lineages. Hence, its presence or absence is “less revealing” than the nature of the male antennule, leg 3 endopod, leg 4 basis, female genital field, and caudal rami in parastenocaridid phylogeny, as opined by Schminke (2010: 358). The genus Proserpinicaris Jakobi, 1972 sensu Karanovic, Cho & Lee, 2012, presently containing 20 species, is Palearctic in distribution, with its centre of diversity lying in southern Europe (Karanovic et al. 2012). Till date, only five Asiatic species of this genus are known: Proserpinicaris nipponicus Chappuis, 1955 from Japan, Proserpinicaris ondali Lee & Chang, 2009, Proserpinicaris imjin Karanovic, Cho & Lee, 2012, Proserpinicaris young Karanovic, Cho & Lee, 2012, and Proserpinicaris wangpi Karanovic, Cho & Lee, 2012, from South Korea. The genus is reported herein from India for the first time. Both Proserpinicaris corgosinhoi n. sp. and Proserpinicaris karanovici n. sp. fulfill all the principal generic criteria, as revised by Karanovic et al. (2012), according to whom the single most important synapomorphic feature of the genus is the presence of a hyaline structure on the anterior surface of the male leg 4 basis, between the exopod and endopod. However, the validity of this character as a synapomorphy of the genus Proserpinicaris has been questioned by Corgosinho et al. (2012a); they argue that what was considered a synapomorphy by Karanovic et al. (2012) is but a symplesiomorphy of the subfamily Forntinalicaridinae Schminke, 2010, and thus renders the genus Proserpinicaris paraphyletic. The genesis of this controversy can be traced to Lang’s (1948) confusing original definition of the proserpinicarisgroup. According to the original definition, this group of species is characterized by the presence of two “appendages” (“Anhangen”) on the male leg 4 basis, but the real difficulty arises in telling apart the endopod from the hyaline process. According to Corgosinho et al. (2012a), while the outer element is the endopod and the inner one the hyaline process in the fontinalicaridinids (e.g. Proserpinicaris proserpina (Chappuis, 1938), Proserpinicaris cantabrica (Chappuis, 1937), and Proserpinicaris phyllura (Kiefer, 1938)), the opposite is the case in certain other parastenocaridids, e. g. Lacustricaris budapestiensis (Török, 1935), etc. We, too, have experienced this problem particularly in the case of P. corgosinhoi n. sp. in which the smooth endopod is small and overlapped by the presumed smooth hyaline process. Although the endopod is similarly overlapped by the hyaline process in P. karanovici n. sp., the former is distinctly larger and ornamented with lateral spinules. Notwithstanding the controversial nature of the key synapomorphy of the genus, the aforesaid two new fontinalicaridinids species fulfill the unique constellation of all other features as given by Karanovic et al. (2012) in the diagnosis of the genus Proserpinicaris. Hence, we thought it advisable to place the two species under Proserpinicaris rather than to assign them to Parastenocaris, the “taxonomic repository”, pending a much deeper analysis of the systematics of the Parastenocarididae as a whole. Proserpinicaris corgosinhoi n. sp. can be easily distinguished from its congeners by the following features: the male antennular second segment, which is generally longest, is only as long as the fifth segment of the same appendage—perhaps a unique character in the Parastenocarididae; the caudal rami are rather short, and the fifth legs have a spiniform endopodal seta (innermost seta) in both sexes; the male leg 3 exopod is stumpy and has a very prominent tubular pore, and the apophysis of the same appendage is mace-like, without apical seta; the male leg 4 has a short foliaceous hyaline structure occurring close to, and partially overlapping, the endopod; and the dorsal window on the preanal segment extends down to the ventral surface in both sexes. Given its unique constellation of the above-mentioned characters, it is hard now to identify the possible sister species of P. corgosinhoi n. sp. among its known congeners. Hence, future investigations on the Indian subcontinent are likely to reveal its closely allied new congeners. Except for both hyaline structure and endopod of the male leg 4 occurring close together, and also the characteristic shape of the former, P. karanovici n. sp. is quite different from P. corgosinhoi n. sp. in all other specific features. Interestingly, however, P. karanovici n. sp. displays a somewhat close morphological kinship not with any of its Asian congeners, but with the Portuguese Proserpinicaris cruzi (Noodt & Galhano, 1969), which is known only by its male (see Noodt & Galhano 1969). The chief resemblances between the two taxa relate to

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similar body size, the male leg 3 having short exopod, longer thumb, and disto-lateral ornamentation, and the male leg 4 endopodal size and ornamentation, caudal rami longer than anal somite and carrying two instead of three lateral setae, and the shape and size of anal operculum. Proserpinicaris karanovici n. sp. can, however, be easily distinguished from P. cruzi by the following features: subproximal part of caudal rami dilated, unornamented vs. cylindrical, ornamented with transverse spinular row; male leg 3 apophysis long vs. short, gradually tapering to acuminate point vs. blunt; endopod represented by strong but short spine vs. long, slender seta; endopod leg 4 rodshaped with distal modified, fused seta vs. lanceolate with serrulate margins; and leg 5 with distinct spiniform process at distal inner corner vs. rounded, with four setae vs. three setae. In India, both the Himalayan and Peninsular River Systems present a vast and ecologically diversified hyporheic realm and riparian areas. In addition, there are over a thousand caves especially in the northeastern Himalayas (Kharpran Daly 2006). This immense groundwater realm, apparently harboring enormous biodiversity, has yet been little investigated for interstitial micro-crustaceans. This is perhaps due to the minute size of the organisms, the difficulties involved in their sampling, the exacting microscopic study and drawing work, the lack of taxonomic expertise and funding support, etc. In our ongoing biodiversity studies of the subterranean groundwater biodiversity of India, we have so far recorded and formally described 13 parastenocaridid species including the three new species that are dealt with here, and also redescribed P. curvispinus, collected mainly in the coastal belt of the rivers Krishna and Godavari and a few caves in peninsular India. Of these 14 species, nine species of the genus Parastenocaris (P. curvispinus, P. gayatri, P. savita, P. mahanadi, P. muvattupuzha, P. sutlej and P. gundlakamma) and one species each of Kinnecaris Jakobi, 1972 (K. godavari) and Siolicaris Jakobi, 1972 (S. sandhya) are from the sandy sediments of the hyporheic zone, two species (Parastenocaris tirupatiensis, Proserpinicaris karanovici n. sp.) are from the phreatic waters of the riparian bore wells, and three species (Parastenocaris kotumsarensis , P. edakkal n. sp., and Proserpinicaris corgosihoi n. sp.) are from interstitial cave pools. The distribution of all these species is as in Fig. 1a. As for the habitat preference of these species, P. curvispinus, P. gayatri and K. godavari, which are generally hyporheic, have also been found in the riparian bore wells. Parastenocaris curvispinus and Parastenocaris mahanadi can tolerate brackish conditions as well (Ranga Reddy & Defaye 2007, and present study). On several occasions, P. gayatri, P. savita, and S. sandhya have been found sympatrically in the hyporheic habitats and also heavily preyed upon by the juveniles of a commercially important gobioid fish, Glossogobius giuris (Hamilton, 1822) ( see Ranga Reddy 2001). The concept of short-range endemism of stygofauna (Harvey 2002; Humphreys 2008; Karanovic & Cooper 2011; and others) is not applicable to P. curvispinus and P. mahanadi, which indeed have a wide-range distribution in the peninsular India. Surprisingly, the Sri Lankan P. curvispinus is most common in the hyporheic habitats of both eastern and western coastal deltaic belts of peninsular India (Ranga Reddy & Defaye 2007). Numerically, parastenocaridids are invariably dominant over bathynellaceans, which are another typical groundwater group. Future stygofaunistic investigations in other parts of India are likely to reveal many more new and interesting taxa. Presently, the hyporheic biodiversity of India, as elsewhere, is under the increasing, disastrous impact of the illegal, rampant sand mining activity of the rivers in the country (see Ranga Reddy 2014). Biogeographically, the Indian Parastenocarididae represent both Gondwanan and Asian elements. While K. godavari and S. sandhya have clear-cut Gondwanan heritage, having their congeners distributed, respectively, in Africa and South America (see Ranga Reddy 2011b), the brevipes-group of Parastenocaris species, with its possible origin in “tropical Asia” (Reid 1995) is represented in India by P. gayatri, P. savita, P. muvattupuzha, P. sutlej, P. gundlakamma, and P. kotumsarensis. Parastenocaris tirupatiensis shares some close affinities with the Neotropical Remaneicaris (see Corgosinho et al. 2012b). Also, two more new Indian taxa that are present in our samples, which display Gondwanan heritage, are under our study. Most of the presently known typical Gondwanan derivatives occur in the peninsular India, “which per se is biogeographically India vera, the largest and the oldest region of the original floras and faunas of India” (Mani 1974).

Acknowledgements We sincerely thank the Department of Science and Technology (DST), Ministry of Science & Technology, New Delhi, for providing the funding support under the Major Project Research Projects (SR/SO/AS/25/2007 and SR/

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SO/AS-21/2011) and also Dr. K. Subhash Babu for his guidance and assistance during our field trip to the remote Edakkal Cave. We must also gratefully acknowledge the helpful comments of the anonymous reviewers and the meticulous editing of the manuscript done by the subject editor.

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