identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
038A87FB192C016598A9FEABE1D8FCB6.text	038A87FB192C016598A9FEABE1D8FCB6.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Bythinia	<div><p>BYTHINIA EVEZARDI W.T. BLANFORD, 1881</p> <p>This species was described by Blanford (1881) from near Khandala at the top of the Bhor Ghat Pass, from material collected by G. Evezard. In the description Blanford (1881) mentioned impressed spiral lines, conical umbilicus, and nearly vertical and angulate aperture with entire peristome. It was made the type of the monotypic bithyniid genus Sataria by Annandale (1920), who added that the calcified operculum had a central nucleus and concentric ridges, and described a bithyniid radula. He also commented that its range was coterminous with that of Cremnoconchus, having been recorded from Mahabaleshwar and Khandala. A collection of juvenile Cremnoconchus from Bhor Ghat in (USNM 317704, ex Evezard Colln) is labelled ‘ Cremnoconchus evezardi Blanford, 1880 ’. This is presumably a misidentification based on the name B. evezardi, rather than a manuscript name for this Cremnoconchus. We are not aware that this taxon has ever been formally assigned to Cremnoconchus.</p> </div>	https://treatment.plazi.org/id/038A87FB192C016598A9FEABE1D8FCB6	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Reid, David G.;Aravind, Neelavara Ananthram;Madhyastha, Neelavara Ananthram	Reid, David G., Aravind, Neelavara Ananthram, Madhyastha, Neelavara Ananthram (2013): A unique radiation of marine littorinid snails in the freshwater streams of the Western Ghats of India: the genus Cremnoconchus W. T. Blanford, 1869 (Gastropoda: Littorinidae). Zoological Journal of the Linnean Society 167 (1): 93-135, DOI: 10.1111/j.1096-3642.2012.00875.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2012.00875.x
038A87FB192C01659885FC0EE025FB66.text	038A87FB192C01659885FC0EE025FB66.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Cremnoconchus W. T. Blanford 1869	<div><p>CREMNOCONCHUS MESSAGERI BAVAY &amp; DAUTZENBERG, 1900</p> <p>This species from Vietnam was initially described in the genus Cremnoconchus by Bavay &amp; Dautzenberg (1900) and was figured under the same name by Dang (1980). Based on our examination of the original figure and description, it is a member of the Paludomidae, similar to Paludomus petrosus (Gould, 1844).</p> </div>	https://treatment.plazi.org/id/038A87FB192C01659885FC0EE025FB66	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Reid, David G.;Aravind, Neelavara Ananthram;Madhyastha, Neelavara Ananthram	Reid, David G., Aravind, Neelavara Ananthram, Madhyastha, Neelavara Ananthram (2013): A unique radiation of marine littorinid snails in the freshwater streams of the Western Ghats of India: the genus Cremnoconchus W. T. Blanford, 1869 (Gastropoda: Littorinidae). Zoological Journal of the Linnean Society 167 (1): 93-135, DOI: 10.1111/j.1096-3642.2012.00875.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2012.00875.x
038A87FB192C0165997EFA8CE153F95B.text	038A87FB192C0165997EFA8CE153F95B.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Lacuninae GRAY 1857	<div><p>SUBFAMILY LACUNINAE GRAY, 1857</p> <p>Risellidae Kesteven, 1903: 621–631. Cremnoconchinae Preston, 1915: 64.</p> <p>Bembiciidae Finlay, 1928: 241.</p> <p>Remarks: Cremnoconchus was first placed in the subfamily Lacuninae by Stoliczka (1868, as Cremnobates). A new subfamily, Cremnoconchinae, was created for this genus, without discussion, by Preston (1915). If it is shown that Cremnoconchus deserves subfamilial or familial rank, the name Cremnoconchinae Preston, 1915 is therefore available.</p> </div>	https://treatment.plazi.org/id/038A87FB192C0165997EFA8CE153F95B	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Reid, David G.;Aravind, Neelavara Ananthram;Madhyastha, Neelavara Ananthram	Reid, David G., Aravind, Neelavara Ananthram, Madhyastha, Neelavara Ananthram (2013): A unique radiation of marine littorinid snails in the freshwater streams of the Western Ghats of India: the genus Cremnoconchus W. T. Blanford, 1869 (Gastropoda: Littorinidae). Zoological Journal of the Linnean Society 167 (1): 93-135, DOI: 10.1111/j.1096-3642.2012.00875.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2012.00875.x
038A87FB192C016A989DF972E0FDFEDE.text	038A87FB192C016A989DF972E0FDFEDE.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Cremnoconchus W. T. Blanford 1869	<div><p>CREMNOCONCHUS W.T. BLANFORD, 1869</p> <p>Cremnobates W.T. Blanford, 1863: 184 (type species by monotypy Cremnobates syhadrensis W.T. Blanford, 1863; not Cremnobates Swainson, 1855; not Cremnobates Günther, 1861).</p> <p>Cremnoconchus W.T. Blanford, 1869: 343 (new name for Cremnobates Blanford, 1863, not Günther, 1861).</p> <p>Littorina (Cremnoconchus) – Stoliczka, 1871: 113.</p> <p>Cremnoconchus (Lissoconchus) Thiele, 1929: 125 (type species by monotypy Cremnoconchus conicus Blanford, 1870).</p> <p>Taxonomic history: Blanford’s (1863) original name Cremnobates was preoccupied ‘for a genus of fishes’ (i.e. Cremnobates Günther, 1861; a name used even earlier by Swainson, 1855) and he therefore renamed it Cremnoconchus (Blanford, 1869). The genus has been treated as valid at generic rank by all subsequent authors except Stoliczka (1871), who suggested that it should be ranked as a subgenus of Littorina. The subgenus Lissoconchus was introduced by Thiele (1929) for C. conicus and has occasionally been employed for this and other Cremnoconchus species without conspicuously ribbed shells (Wenz, 1938; Subba Rao &amp; Mitra, 1979; Subba Rao, 1989; Ramakrishna &amp; Dey, 2007).</p> <p>Diagnosis: Shell: turbinate; smooth or with spiral ribs; protoconch smooth, less than 1.5 whorls (indicating non-planktotrophic development). Operculum: paucispiral; calcified, with proteinaceous layer internally and externally. Male: prostate closed; prostate gland subepithelial; anterior vas deferens closed; penial vas deferens (sperm duct) deeply closed (i.e. no epithelial connection to surface); penial filament retracted into cavity in base; simple penial glands in base, sometimes forming a short protruberance. Female: seminal receptacle absent; oviducal glands subepithelial; egg groove coiled in one spiral. Radula: 5 cusps (plus 2 denticles) on rachidian; 3–5 cusps on outer marginal. Alimentary system: salivary glands anterior to nerve ring. Nervous system: pleurosuboesophageal connective short. (Slightly modified from Reid, 1989.)</p> <p>Shell: The shell shape lies within the range of that of marine littorinids, being globular, turbinate or highturbinate, of moderate thickness, and sometimes with strong spiral ribs (Figs 3, 9, 11). As in many freshwater molluscs, the periostracum is relatively thick. Dissolution of the shell, or radulation by other snails, occurs where the periostracum is damaged, and the apical whorls are frequently eroded away. An umbilicus is present in some species, but is usually absent in juveniles, so it is correctly termed a ‘pseudumbilicus’. The surface is marked by prominent, fine or indistinct spiral microstriae that are visible under low magnification (Fig. 4) and are a useful feature for identification. As in most marine littorinids (e.g. Reid, 1986a, 1996, 2007) the shell shows striking intraspe-</p> <p>N, sample size; H2/B, shell shape index; H2/LA, relative spire height index; LA/WA, apertural shape index; DS/OL, opercular ratio; SE, standard error of the mean. See Figure 1 for definition of dimensions B, DS, H, H2, LA,OL, and WA.</p> <p>cific variability. In most species the sampling remains too limited to observe patterns in the shell variation. However, in the most well-studied species, C. syhadrensis, there are clear differences between the range of variation within each of four local populations (Tables 1 and 2), suggesting possible genetic differentiation. In one population (from Torna; Fig. 3F, J–M) the shells range from almost smooth to strongly ribbed, so that development of ribs is not a reliable character for identification. Similarly, in C. canaliculatus a sutural rib can be present or absent (Fig. 9P– EE). In both C. conicus and C. canaliculatus the overall shape of the shell (relative spire height and peripheral angulation) and development of the pseudumbilicus are variable (Fig. 9). Shell coloration ranges from pale to dark brown in these three relatively well-sampled species, sometimes with 1–3 indistinct brown bands (Figs 3, 9), but does not appear to be polymorphic (i.e. with discrete morphs) within populations. There is slight sexual dimorphism in size, males being smaller (as also noted by Stoliczka, 1871); this phenomenon is usual in littorinids (Reid, 1986a, 1996). The protoconch is similar in all species in which it has been seen (Fig. 16C–F); it is smooth, of 1.3–1.5 whorls, 0.47–0.70 mm in diameter, and therefore similar to the protoconchs of marine littorinids with non-planktotrophic development (Reid, 1989).</p> <p>Headfoot and locomotion: Externally, the animal is typical of the family, with blunt snout and mobile cephalic tentacles with eyes at their bases (Fig. 2H, I). The head and sides of the foot are commonly pigmented grey to black, but the animal is paler in C. canaliculatus, and in C. syhadrensis some populations are virtually unpigmented. Where head pigmentation is pale, the reddish myoglobin of the buccal mass musculature is visible by transparency (Fig. 2I). The sole of the foot does not show an anterior transverse (propodial) groove, but a small anterior pedal gland is present (Reid, 1989). No longitudinal division of the sole is visible externally, but locomotion is nevertheless by ditaxic retrograde waves of muscular contraction; one complete wave passes back on one side of the sole of the foot, then on the other.</p> <p>Operculum: The operculum of Cremnoconchus is paucispiral, but thickened by calcification in between internal and external proteinaceous layers. In species with the thickest calcification (e.g. C. syhadrensis) the calcified part is inflexible, but an uncalcified flange remains around the margin. Internally, most species show a thickened spiral boss or rib of organic material; this, however, does not show well in SEM images because it is of relatively low relief (Fig. 5). The tightness of coiling, degree of calcification and development of the internal ridge show some variation between species (Table 1; Fig. 5). The calcification of the operculum was mentioned by several early authors (Blanford, 1863; Stoliczka, 1871), but without further comment. In fact such calcification is almost unique among littorinids (Reid, 1989), the only other case being the aragonitic calcareous deposit on the outer surface of the operculum of Tectarius niuensis</p> <p>or for</p> <p>grey</p> <p>,</p> <p>but tencles snout</p> <p>colour dark</p> <p>Animal Pale to black Unpigmented pale grey tip of and??</p> <p>faint</p> <p>columella red-brown bands; aperture</p> <p>and suture</p> <p>at;; darker columella</p> <p>spire</p> <p>darker</p> <p>white;</p> <p>bands red-brown bands; usually and</p> <p>colour</p> <p>Shell Pale</p> <p>yellow-brown</p> <p>;</p> <p>; bands</p> <p>aperture</p> <p>brown stained purple-brown with broad Mid-brown spire; aperture with columella white yellow-brown no, Pale columella white; Pale yellow-brown on; aperture and spire white</p> <p>Umbilicus mm () – 0.6 1.2 0.2 –0.3 0.3 0.1 – 0.1 –0.5</p> <p>0</p> <p>ribs ribs are</p> <p>3, as, 0 as, 2 – ribs</p> <p>syhadrensis Base – 24 threads 15 developed. 20 threads c developed threads 5 10 – innermost ribs; threads 0 0 Cremnoconchus of Ribs above periphery – 10 8 8 8 5 – at shoulder (2 and; periphery) to or up 8 populations. Max shell height) (mm 9.4 8.3 5.9 9.8</p> <p>C I E</p> <p>of, –,</p> <p>Comparison</p> <p>.</p> <p>Shell figures. 3 Fig</p> <p>B Fig. G 3 Fig. D 3</p> <p>Hill</p> <p>, F. Fig 3</p> <p>120</p> <p>M – J)</p> <p>2 Table Locality (sample) size N Matheran () 200 Khandala () 200, Anjani Nashik 10) (Torna Fort (</p> <p>(Reid &amp; Geller, 1997). The operculum can be withdrawn to a depth of about 0.1–0.2 of the final whorl, but up to 0.4 in C. agumbensis.</p> <p>Male reproductive system: The anatomy and histology of the reproductive system have been described by Linke (1935) and Reid (1989). A closed prostate with subepithelial glands leads to a tubular anterior vas deferens and to the cephalic penis on the right side of the headfoot. The littorinid penis often bears elaborate glandular structures, but in Cremnoconchus the subepithelial, neutral mucous glands (staining blue in trichrome) are usually simply scattered in parts of the penial base, which appear swollen and opaque. In C. conicus and C. canaliculatus the subepithelial glands are found in a basal drum-shaped appendage, but this is unlike the penial glandular disc or complex mamilliform penial glands present in some marine littorinids (Reid, 1989). A feature of Cremnoconchus that is unique in the family is the invagination of the penial filament (the intromittent part) into a distal pocket of the muscular base. The filament can be extended to a considerable length and in preserved material is usually fixed in a partly extended state. In the living animal, however, the filament is generally retracted. Dissection reveals that the penial vas deferens (sperm duct) within the base is coiled and folded to allow for extension of the filament during copulation (the duct in Fig. 6F, G is 2.5 times the length of the penial base), and lies within an internal space that is presumably involved in the hydrostatic extension of the filament. Withdrawal of the filament is accomplished by a retractor muscle lying adjacent to the coiled vas deferens and inserted at the junction of the base of the filament with the bottom of the invagination (Fig. 6G). Before entering the filament, the vas deferens becomes tightly bound to the retractor muscle. As in most littorinid genera, the form of the penis is an important character for discrimination</p> <p>› Figure 2. Habitats and living animals of Cremnoconchus species. A, 10 km west of Mahabaleshwar on road to Poladpur, Raigad Dist., Maharashtra; altitude 692 m (C. canaliculatus). B, 6 km west of Mahabaleshwar on road to Poladpur; 1103 m (C. conicus). C, Hulikal Ghat, Udupi Dist., Karnataka; 475 m (C. cingulatus, C. dwarakii). D, escarpment of Western Ghats at Mahabaleshwar, Maharashtra; this stream is now polluted and contains no Cremnoconchus. E, C. globulus in flowing water at Lesser Kadambi Falls, Chikmagalur Dist., Karnataka; 967 m. F, Khandala, Pune Dist., Maharashtra; 297 m (C. syhadrensis). G, C. syhadrensis aestivating under ledges at Khandala; 297 m. H, living C. conicus, 6 km west of Mahabaleshwar on road to Poladpur. I, living C. syhadrensis, Khandala.</p> <p>of species and may be a component of the specific mate-recognition system (Reid, 1996).</p> <p>Spermatozoa: These have been observed only in C. castanea, by light microscopy. They consist only of euspermatozoa (i.e. parasperm are absent), which are filiform and up to 193 Mm in length. These are among the longest recorded for Littorinidae, exceeded only by those of Bembicium (Reid, 1989). The sperm head is 24 Mm long and a distinct helical structure is visible. This may correspond to the helical condensation of chromatin in the nucleus of developing spermatids, reported in SEM studies of sperm development in Littorina species (Buckland-Nicks &amp; Chia, 1976; Paviour, Mill &amp; Grahame, 1989); however, these reports have not mentioned a helical structure in mature eusperm.</p> <p>Female reproductive system: The spiral route of the egg groove through the pallial oviduct is a well-known synapomorphy of Littorinidae (Reid, 1989). In Cremnoconchus the egg groove forms a single spiral loop of about 3.5 revolutions. Linke (1935) discriminated albumen, capsule and shell glands in the pallial oviduct, but did not explain the spiral structure. Histological examination suggests that the subepithelial glandular material composing the spiral portion of the pallial oviduct is mainly albumen gland, differentiated into two parts staining darker and paler blue in trichrome; these parts can sometimes be distinguished as more and less opaque cream areas in preserved specimens (Fig. 6C). The subepithelial lining of the final straight section of the pallial oviduct stains reddish in trichrome and appears to be responsible for the secretion of the outer coating of the egg capsule, which stains similarly; this gland is not, however, homologous with the capsule gland of the Littorininae (Reid, 1989). The ciliated cells lining the egg groove do not contain black pigment. There is a large copulatory bursa, opening in an anterior position (Fig. 14C). The seminal receptacle typical of most littorinids is absent; sperm are instead stored in the renal oviduct (Linke, 1935; Reid, 1989).</p> <p>Eggs: Mature ova can sometimes be seen in the distal part of the ovarian oviduct. In the pallial oviduct these are coated with albumen and a firm gelatinous covering, to attain a final diameter usually in the range 0.40–0.49 mm. Between 1 and 5 such eggs can be found in the straight section of the pallial oviduct (Figs 6C, 15J); these are either uncleaved or show only the first cleavage of the embryo, suggesting that they are laid before further development proceeds.</p> <p>Radula: The radula of Cremnoconchus has been drawn by Troschel (1867), Stoliczka (1871), Annan- dale &amp; Prashad (1919) and Reid (1989), but SEM images are given here for the first time (Figs 7, 13, 16A, B). The radula is long (length relative to shell height 0.82–4.24) and the radular sac is coiled over the mid-oesophagus. As in all littorinids it is taenioglossate, with all seven teeth in each row well developed and a ‘littorinid notch’ in the base of the lateral tooth (Reid, 1989). The base of the rachidian tooth is relatively narrow, lacking lateral projections (Troschel, 1867), but otherwise the radula is of a generalized littorinid form (Reid, 1989).</p> <p>Mantle cavity: When the anatomy was first described, Cremnoconchus was said to possess a vascular mantle cavity (a ‘pulmoniferous sac’) with no trace of gills (Blanford, 1863). Gills were subsequently described (Stoliczka, 1871; Annandale, 1919; Prashad, 1925). The leaflets are of low triangular outline, continued as ridges (not branched as stated by Stoliczka, 1871) across the roof of the mantle cavity, and number 23–50. Although it has been reported that the opening of the mantle cavity is small and can be completely closed (Annandale, 1919; Annandale &amp; Prashad, 1919), in fact the mantle edge and wide mantle opening are the same as in marine littorinids. The observation may be a reference to the fact that, when active, the headfoot largely occludes the shell aperture. The osphradium is short (Prashad, 1925), only about one-third of the extent of the row of gill leaflets, but it is not ‘papilliform’ (Annandale, 1919; Annandale &amp; Prashad, 1919). The hypobranchial gland is vestigial (Prashad, 1925).</p> <p>Range: Endemic to the northern and central Western Ghats of India, in the states of Maharashtra and Karnataka (Fig. 8).</p> <p>Habitat and ecology: The habitat is highly specific, restricted to streams and waterfalls on the basaltic cliffs of the western escarpment of the Western Ghats (Fig. 2D), between altitudes of 297 and 1125 m (and potentially up to 1400 m, from localities of museum specimens; Supporting Table S1; see also Blanford, 1863, 1870; Annandale, 1919; Prashad, 1925). During the heavy rain of the south-western monsoon season (June–September) these streams become torrents. However, when collections were made by the authors during October 2010 the snails were mostly to be found crawling over rocks wetted by spray, or where water was flowing in a shallow film, usually in abundance (Fig. 2E). Cremnoconchus conicus also occurred in shallow pools with moving water, while C. syhadrensis and C. dwarakii were sometimes found on moss and wet vegetation beside the streams. In the northern part of the range (Maharashtra state) the surrounding vegetation in the stream gullies consisted of fern and balsam (Impatiens), with adjacent woodland or cloud forest, depending upon altitude (Fig. 2A, B). In the southern part (Karnataka state) the streams ran through rainforest (Fig. 2C). During the dry season, small streams may disappear and C. syhadrensis is known to aestivate in clusters in crevices and shaded pits in the rock (Blanford, 1863; Annandale, 1919; Hora, 1926, 1928). At Khandala (October 2010) C. syhadrensis was observed in tight clusters of up to 1000 individuals under overhangs on shaded cliffs, where they would presumably aestivate as the rock surface dried out (Fig. 2G).</p> <p>The behaviour of C. syhadrensis at Khandala was described by Annandale (1919: 119). At this locality it was abundant on cliffs, shaded from the midday sun, which ‘supported a growth of the peculiar dull green filamentous alga on which it feeds’. The animals became inactive and closed the operculum tightly when dry. When submerged, the snails crawled out of the water but, if prevented from reaching the surface, drowned after 24 h. The mantle cavity was said to be filled with water (‘never filled with air’) in snails that were crawling in air, but this was not the case in animals observed during the present study, in which the mantle cavity was air-filled. Stoliczka (1871) noted that the stomach of this species contained ‘minute algae’. In the present study the faecal pellets of C. hanumani were found to consist almost entirely of elongated diatoms.</p> <p>The breeding season has not been recorded, but is presumed to be during the monsoon season. In October 2010, at the end of the monsoon, some specimens of all the species collected were mature (i.e. seminal vesicle with sperm and/or eggs in ovarian or pallial oviduct), but some were spent, suggesting that the breeding season was coming to an end. Samples of C. castanea collected in March and June also included some mature individuals (BMNH). Stoliczka (1871) recorded eggs in a mature female of C. syhadrensis in March. Since the eggs in the final section of the oviduct were always either uncleaved or at the twocelled embryonic stage, it is predicted that the large eggs are deposited and not retained until hatching. There is no evidence for ovoviviparity, as occurs in some marine littorinids (Reid, 1989). Eggs are probably laid either singly or in small numbers, and presumably on the rock surface, where development and hatching take place.</p> <p>The prevalence of parasitism by trematodes was very low, being observed only in C. canaliculatus from a single locality.</p> <p>Remarks: At present there is no strong evidence for the subgeneric division of Cremnoconchus. The recognition of Lissoconchus as a subgenus based only on the smooth shell of C. conicus (Thiele, 1929) is unwarranted, in view of the variability of shell ribs in C. syhadrensis.</p> </div>	https://treatment.plazi.org/id/038A87FB192C016A989DF972E0FDFEDE	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Reid, David G.;Aravind, Neelavara Ananthram;Madhyastha, Neelavara Ananthram	Reid, David G., Aravind, Neelavara Ananthram, Madhyastha, Neelavara Ananthram (2013): A unique radiation of marine littorinid snails in the freshwater streams of the Western Ghats of India: the genus Cremnoconchus W. T. Blanford, 1869 (Gastropoda: Littorinidae). Zoological Journal of the Linnean Society 167 (1): 93-135, DOI: 10.1111/j.1096-3642.2012.00875.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2012.00875.x
038A87FB192301779BD6FEF5E67BFC56.text	038A87FB192301779BD6FEF5E67BFC56.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Cremnoconchus syhadrensis (W. T. Blanford 1863)	<div><p>CREMNOCONCHUS SYHADRENSIS</p> <p>(W.T. BLANFORD, 1863)</p> <p>(FIGS 2G, I, 3, 4B, C, 5A, B, 6, 7A, B, 8, 16C)</p> <p>Cremnobates syhadrensis Blanford, 1863: 184–186, pl. 4, figs 1–7 (fig. 6, crawling animal; figs 4, 5, operculum) (Western Ghats, in the neighbourhood of Bombay, India; restricted to Khandala by Subba Rao &amp; Mitra, 1979; here corrected to Matheran, Raigad District, Maharashtra; neotype, here designated, BMNH 1906.1.1.2247, Fig. 3A). Troschel, 1867: 90–94, pl. 2, fig. a (radula).</p> <p>Cremnoconchus syhadrensis – Blanford, 1869: 343. Stoliczka, 1871: 108–115, figs 1–3 (anatomy), 4, 5 (radula). Hanley &amp; Theobald, 1876: 58, pl. 146, fig. 6. Theobald, 1876: 12. Nevill, 1885: 173. Tryon, 1887: 256, pl. 46, fig. 47. Preston, 1915: 65 (as syhadensis). Annandale &amp; Prashad, 1919: 149, pl. 4, figs 2, 3 (anatomy), 4 (radula) (in part, includes C. canaliculatus). Prashad, 1925: 138–139, fig. 2C (mantle cavity). Linke, 1935: 72–87, figs 1–8 (anatomy). Reid, 1989: figs 1i, 4c, d (operculum), 6i (penis), 9i (pallial oviduct). Aravind et al., 2011: 58 (unnumbered fig.) (BMNH 1906.1.1.2247, same lot as neotype of C. syhadrensis).</p> <p>Cremnoconchus (Cremnoconchus) syhadrensis – Thiele, 1929: 125, fig. 98. Wenz, 1938: 522, fig. 1376. Subba Rao &amp; Mitra, 1979: 24–25. Subba Rao, 1989: 63, figs 88, 89, 89a, b. Ramakrishna &amp; Dey, 2007: 111, fig. 67A, B.</p> <p>Cremnoconchus ‘ from Mahabaleshwar’ Aravind et al., 2011: 58, unnumbered fig. (locality in error; BMNH 1906.1.1.2247, same lot as neotype).</p> <p>Taxonomic history: No types were segregated in BMNH, but there exists a sample of four specimens from W. T. Blanford’s collection labelled, in his hand, ‘ Cremnobates syhadrensis picked specimens’, but with no locality. One of these is here designated neotype (Fig. 3A). The type locality was restricted to Khandala by Subba Rao &amp; Mitra (1979), but this is here corrected to Matheran (see Remarks). The specific name is derived from ‘Sahyadri’ (erroneously spelled ‘Syhadri’ by Blanford, 1863), the name for the Western Ghats in Marathi and other Indian languages.</p> <p>Diagnosis: Shell turbinate with 2–8 spiral ribs above periphery; pseudumbilicus present or absent; surface dull, with fine or indistinct microstriae. Operculum thickly calcified, with internal ridge. Penis with distal glandular swelling of base, no lateral glandular</p> <p>flange or drum-shaped gland, stout filament. Western Maharashtra State.</p> <p>Shell (Figs 3, 4B, C): Shell H 5.0– 9.9 mm. Shape (Table 1) broadly turbinate; whorls rounded, weakly angled at periphery; suture distinct; apex intact or eroded. Columella moderately broad. Pseudumbilicus usually conspicuous and perforated (Fig. 3F), 0.6– 1.2 mm wide, with sharply angled margin; sometimes narrow and not perforated (Fig. 3H), 0.1–0.5 mm; may be closed in juveniles. Upper surface: usually 5–10 (mode 8) narrow ribs at and above periphery, sometimes becoming weak on last whorl (Fig. 3J); ribs (especially on spire) may be slightly granulated by intersection with growth lines; 0–3 threads in interspaces between grooves; shell sometimes smooth but for narrow rib at shoulder (giving broad sutural ramp and turreted spire profile) and at angled periphery (Fig. 3L, M). Base: usually 15–24 threads, becoming finer and ultimately indistinct towards pseudumbilicus (Fig. 3C); sometimes 5–10 ribs, of which innermost 2–3 develop as narrow ribs (Fig. 3E); base may lack sculpture (Fig. 3F). Entire surface dull, covered with fine or indistinct spiral microstriae (Fig. 4B, C). Protoconch (Fig. 16C) 1.3–1.4 whorls; diameter 0.47– 0.60 mm; diameter of first whorl 0.29–0.45 mm. Colour: pale yellow-brown, often with red-brown spire (shell fading to white with purplish or pinkish spire; Fig. 3A); some have narrow brown band at suture and broader brown band between shoulder and periphery (Fig. 3G, I); base sometimes with broad brown band but paler around umbilicus (Fig. 3H); columella white, columella and umbilicus sometimes stained purple-brown (Fig. 3C); aperture yellowish with redbrown bands showing through.</p> <p>Animal: Head, tentacles, and sides of foot pale to dark grey or black; sometimes unpigmented except for grey tentacles and tip of snout (Fig. 2I). Gills: 30–40 leaflets; unpigmented. Operculum (Table 1; Fig. 5A, B): opercular ratio high, 0.471 –0.600; strongly calcified, with thickened internal ridge. Penis (Fig. 6A, B, D–G): unpigmented; base wrinkled, distal 20–60% smooth, swollen, opaque and glandular, invagination 20–40% of length of base in ethanol-fixed specimens; filament relatively stout, usually protruding in ethanol-fixed specimens, tip can be retracted to about half length of penial base in living animals. Pallial oviduct (Fig. 6C): as for genus. Pallial oviduct containing 1– 5 eggs 0.24–0.49 mm diameter, including transparent covering 0.06 mm thick (observed alive).</p> <p>Radula (Fig. 7A, B): Relative radula length 1.60–4.24. Rachidian: length/width 1.51–1.57; 5 cusps (+ 1 outer denticle on either side). Lateral: 5 cusps (+ 1 inner denticle). Inner marginal: 5 cusps (+ 1 outer denticle; sometimes + 1 inner denticle). Outer marginal: 3–5 cusps (sometimes + 1 outer denticle). Major cusp of each of 5 central teeth in each row elongate leafshaped with rounded tip; other cusps pointed.</p> <p>Range (Fig. 8): Western Maharashtra State, from about 30 km west of Nashik to 50 km south-west of Pune (185 km linear distance). Records (Supporting Table S1): Maharashtra State: Anjani, near Trimbakeshwar, Nashik District (BMNH 1907.9.25.3–4); Igatpuri (Blanford, 1870; as Egutpoora); Matheran (BMNH 20120035; ZSI/WGRS/IR.INV-2290, 2291); Bhor Ghat (BMNH 1868.12.11.38); Khandala (BMNH 1888.12.4.823–826; ZSI/WGRS/IR.INV-2292, 2293); Karlee Hill Fort, Kurkulla, near Pune (locality tentatively identified as Karli; USNM 317697); Vadgaon, near Pune (USNM 317687); Sinhgarh Fort, near Pune (USNM 317691); Torna Fort, near Pune (USNM 317695, 317696, 317683); Rajgarh Fort, near Pune (USNM 317688).</p> <p>Habitat and ecology: On unshaded east-facing cliff with water seeping over rock; 442 m altitude; common, found individually, with many juveniles (Matheran population; 17 Oct. 2010). On wet basalt cliffs, clustered in tightly packed groups of up to 1000 under overhangs 30 cm in width; 297 m altitude (Khandala population; 17 Oct. 2010; Fig. 2F, G). Hora (1926: 448) recorded it as ‘quite plentiful on big rocks projecting out of the water at the edge of the pool in the neighbourhood of the fall [near Khandala, at end of December]. They were found aestivating in small pits well protected from the midday sun.’ Annandale (1919) also observed it in great abundance on cliffs at Khandala, shaded from the midday sun, where animals were active in spray from the falls, feeding on a ‘dull green filamentous alga’. Stoliczka (1871) noted that the stomach contained ‘minute algae’. Collected from 4000 ft (1219 m) at Anjani (BMNH 1907.9.25.3– 4). Recorded localities range up to 1400 m (Supporting Table S1).</p> <p>Remarks: This is the most well-known species in the genus and has been the subject of several anatomical studies (Stoliczka, 1871; Annandale &amp; Prashad, 1919; Prashad, 1925; Linke, 1935; Reid, 1989) and ecological notes (Stoliczka, 1871; Annandale, 1919; Hora, 1926). It was found to be common at Matheran and extremely abundant at Khandala (October 2010). However, Matheran is a tourist destination where development is proceeding and Khandala lies in the Bhor Ghat Pass, where motorways are under construction to carry heavy traffic between Mumbai and Pune. The future of these sites cannot be regarded as secure. The Matheran site is the only known locality at which Cremnoconchus occurs outside the defined limits of the Western Ghats region (Fig. 8).</p> <p>Populations of this species show considerable differentiation in shell form across the range. The type locality was given only as ‘in the neighbourhood of Bombay [Mumbai]’ in the original description by Blanford (1863: 184), but he subsequently recorded it from ‘Khandalla [Khandala] where the first specimens were obtained, but also on Matheran hill and at Egutpoora [Igatpuri]’ (Blanford, 1870: 11). His description was of a spirally ribbed, umbilicate, white shell with reddish apex. This, and the (unlocalized) collection from which the neotype (Fig. 3A) has been selected, correspond most closely with modern samples from Matheran (Fig. 3B, C). Differences among available local populations are summarized in Table 2. There is variation in adult size, development of ribs both above and below the periphery, size of umbilicus and colour. In USNM there are three large</p> <p>› Figure 5. Opercula of Cremnoconchus species. A, B, C. syhadrensis, Matheran, Raigad Dist., Maharashtra. C, D, C. dwarakii, Hulikal Ghat, Udupi Dist., Karnataka. E, F, C. conicus, 44 km west of Bhor, Pune Dist., Maharashtra. G, H, C. canaliculatus, 5 km south of Mahabaleshwar, Satara Dist., Maharashtra. I, J, C. hanumani, Hanuman Gundi Falls, Chikmagalur Dist., Karnataka. K, L, C. globulus, Lesser Kadambi Falls, Chikmagalur Dist., Karnataka. M, N, C. agumbensis, Agumbe, Udupi Dist., Karnataka. O, P, C. cingulatus, Hulikal Ghat, Udupi Dist., Karnataka. Q, R, C. castanea, Arasinagundi Falls, Udupi Dist., Karnataka. Abbreviation: b, thickened boss on inner surface of operculum of C. syhadrensis (indistinctly visible by SEM because of low relief and therefore indicated with dotted outline).</p> <p>samples from Torna Fort, near Pune, from the Evezard Collection, made in the late 19th century. These were originally sorted and labelled according to their sculpture: ‘ C. syhadrensis ’ (normal eightribbed shells; USNM 317683), ‘ C. conicus var. canal- iculatus ’ (mostly two ribs, at shoulder and periphery only; USNM 317696); ‘ C. conicus var.’ (two-ribbed shells, but mixed with some C. conicus and C. canaliculatus; USNM 317695). Intermediate forms link the extremes of eight- and two-ribbed shells (Fig. 3F, J–M) and these three samples are suggested to be from a single very variable population. The opercula of the Torna samples are the same as those of specimens from other localities, with strong calcification and a prominent inner ridge, but no anatomical material from Torna was available.</p> <p>There is also variation in the size of eggs and in the diameter of the first whorl of the protoconch. In a female from Matheran the three eggs in the pallial oviduct ranged from 0.24 to 0.39 mm in diameter and the diameter of the first whorl (N = 3) from 0.29 to 0.39 mm in the few shells with intact apex in the sample. The overall ranges of these parameters in the species were larger still (see description above).</p> <p>The shell of C. syhadrensis is the most robust of the Cremnoconchus species, being relatively thick and reinforced by spiral ribs, and the operculum is more thickly calcified than in the others. It is not known if this reflects selection by crushing predators or other mechanical forces. The frequency of repaired shell breakages (remaining as scars on the shell) has been used as an index of sublethal predation or damage (e.g. Reid, 1992). In a total of 208 adult shells (of the Matheran and Khandala types; Table 2) in BMNH, 20 (9.6%) showed a major repaired breakage. The cause is not known, but this is a low level of sublethal damage in comparison with that seen in some marine littorinids (Reid, 1992).</p> <p>Only in one sample from Torna (USNM 317695) is this species mixed with others, C. conicus and canaliculatus, implying that all three are found there together. There are also separate lots of C. syhadrensis and C. canaliculatus from both Bhor Ghat and Karlee. Annandale &amp; Prashad (1919: 149) remarked that ‘ two types of shells occur in a large series from the cliffs at Khandalla. The commoner of these agrees well with Blanford’s figure [i.e. Blanford, 1863: figs 1–7], but in a few specimens the upper surface of the body-whorl is flattened and grooved much as in the same author’s var. canaliculatus of C. conicus.’ This is interpreted as a reference to C. canaliculatus occurring with C. syhadrensis, although in the present study only C. syhadrensis was encountered at Khandala in the Bhor Ghat Pass. At these localties of likely sympatry the shell characters of each species remain distinct, supporting their separate species status.</p> <p>The low spire, 2–8 ribs, weak microstriae and strongly calcified operculum are the diagnostic traits of C. syhadrensis (see Table 3 for distinction of the three sympatric species). Confusion is possible with C. canaliculatus, but this never develops a distinct peripheral rib, is sometimes tall-spired, has stronger microstriae and a less thickly calcified operculum.</p> <p>CREMNOCONCHUS CONICUS W.T. BLANFORD, 1870</p> <p>(FIGS 2H, 4D, E, 5E, F, 7C, D, 8, 9A–O, 10A–H)</p> <p>Anculotus carinatus Layard, 1855: 94 (Mahakeshwar Hills [Mahabaleshwar], Bombay Presidency, India; lectotype, here designated, plus 4 paralectotypes BMNH 20120031, Fig. 9D, E; not Anculotus carinatus Anthony 1840).</p> <p>Cremnoconchus carinatus – Blanford, 1869: 343. Blanford, 1870: 11–12, pl. 3, fig. 5. Stoliczka, 1871: 109, 112 (in part, includes C. canaliculatus). Theobald, 1876: 13. Nevill, 1885: 172. Preston, 1915: 66. Aravind et al., 2011: 58, unnumbered fig. (BMNH 20120031, lectotype of A. carinatus).</p> <p>Cremnoconchus (Lissoconchus) carinatus – Subba Rao, 1989: 63, figs 90, 91. Ramakrishna &amp; Dey, 2007: 112, fig. 68A, B.</p> <p>Anculotus carinatus ‘Anthony’ – Reeve, 1860: Anculotus sp. 42, pl. 5, fig. 42 (not Anthony, 1840; figure is one of the types BMNH 20120031 of A. carinatus Layard, 1855).</p> <p>Cremnoconchus conicus W.T. Blanford, 1870: 10–12, pl. 3, fig. 3 (Torna, 35 miles W of Poona, India; lectotype, here designated, BMNH 1906.1.1.2239, Fig. 9B; 17 probable paralectotypes BMNH 1906.1.1.2242). Hanley &amp; Theobald, 1876: 58, pl. 146, figs 8, 9. Theobald, 1876: 13. Nevill, 1885: 172. Tryon, 1887: 256, pl. 46, fig. 44 (in part, includes C. canaliculatus). Preston, 1915: 65–66. Aravind et al., 2011: 58, unnumbered fig. (BMNH 1906.1.1. 2239, lectotype of C. conicus).</p> <p>Cremnoconchus (Lissoconchus) conicus – Thiele, 1929: 125. Wenz, 1938: 522, fig. 1377. Subba Rao &amp; Mitra, 1979: 25. Subba Rao, 1989: 64, figs 92, 93. Ramakrishna &amp; Dey, 2007: 112–113, fig. 69A, B.</p> <p>Cremnoconchus conicus var. edecollata Nevill, 1885: 172 (near Poona; 12 syntypes ZSI, not seen). Tryon, 1887: 256.</p> <p>Cremnoconchus (Lissoconchus) conicus var. edecollata – Subba Rao &amp; Mitra, 1979: 25–26.</p> <p>Cremnoconchus carinatus var. gigantea Nevill, 1885: 173 (Bombay Presidency; 3 syntypes probably in ZSI, not seen). Tryon, 1887: 256.</p> <p>Taxonomic history: This was the first of the Cremnoconchus species to be discovered, and was originally described as a species of Anculotus (an incorrect subsequent spelling of Anculosa and synonym of Leptoxis, Pleuroceridae; E. Strong, pers. comm.). This grouping among freshwater cerithioideans is explained by the resemblance of the shell to those of globose pleurocerids, including the North American Leptoxis species. Layard’s name Anculotus carinatus is a junior primary homonym of that of Anthony (1840), introduced for a genuine North American pleurocerid. Owing to the confusion of these homonyms, in his monograph of Anculotus Reeve (1860)</p> <p>K</p> <p>I</p> <p>4</p> <p>–</p> <p>)</p> <p>ridge deep</p> <p>EE</p> <p>)</p> <p>–</p> <p>9</p> <p>P</p> <p>perforated Fig. (internal small; at base; filament</p> <p>tall.</p> <p>Fig</p> <p>()</p> <p>wide</p> <p>and</p> <p>microstriae calcified,</p> <p>canaliculatus to Globular shoulder 1 or (Narrow, or Absent strong; Moderately.) G H 5 Fig, Bluntly</p> <p>base</p> <p>tapering drum-shaped</p> <p>gland</p> <p>invagination</p> <p>slender</p> <p>;.</p> <p>C</p> <p>0 Dull (10 N – Fig I (.)</p> <p>Maharashtra) microstriae concave ridge, swollen; slender western</p> <p>in</p> <p>– A</p> <p>Fig</p> <p>.</p> <p>(</p> <p>)</p> <p>9</p> <p>O</p> <p>at</p> <p>rib</p> <p>shoulder perforated</p> <p>never or</p> <p>indistinct internal</p> <p>, no base</p> <p>slightly drum-shaped</p> <p>deep</p> <p>invagination</p> <p>A</p> <p>)</p> <p>10 –</p> <p>H</p> <p>canaliculatus to tall 1 slight absent, sheen; fine 4 E,) D calcified) F 5 E, large;; large; base at Fig. (. C</p> <p>and C.</p> <p>conicus Turbinate 0 (rarely Wide Usually Satin (. Fig Weakly Fig (. Relatively distally gland filament</p> <p>conicus</p> <p>,</p> <p>.</p> <p>C</p> <p>absent A</p> <p>,</p> <p>. Fig 5; no; stout</p> <p>syhadrensis 8) to rarely, microstriae (ridge swelling invagination) D – G Cremnoconchus</p> <p>syhadrensis</p> <p>(</p> <p>. Fig 3) and periphery and perforated</p> <p>wide</p> <p>indistinct</p> <p>or</p> <p>C)</p> <p>B,</p> <p>, internal calcified distal glandular small; gland B, Fig A 6,. (</p> <p>of features. C Turbinate (2 shoulder Moderate Often Dull; fine. Fig (4 Thickly) B with Base lateral filament</p> <p>Diagnostic.</p> <p>Table</p> <p>3</p> <p>Character Shell shape ribs Shell Columella Pseudumbilicus Shell surface Operculum Penis</p> <p>illustrated one of the syntypes of Layard’s species as an example of that of Anthony and gave a North American locality. The shells of the two are not similar.</p> <p>When Blanford (1870) described the species he considered that its range of variability included shells with a strong shoulder rib, named var. canaliculatus, here shown to be a distinct species. There has been persistent confusion in the taxonomic literature over the names conicus, carinatus, canaliculatus and fairbanki, owing to misguided attempts to classify the two species involved, C. conicus and C. canaliculatus, using shell shape and prominence of spiral ribs. Both these characters are variable within species (see Remarks below).</p> <p>The shells described by Nevill (1885) as var. edecollata are a sample of the nominal species with intact spires (fide Subba Rao &amp; Mitra, 1979, who examined the syntypes).</p> <p>Diagnosis: Shell turbinate to tall, lacking ribs; pseudumbilicus absent; surface with satin sheen, microstriae fine or indistinct. Operculum weakly calcified, without internal ridge. Penis with base</p> <p>› Figure 9. Shells of Cremnoconchus species. A –O, C. conicus. P–EE, C. canaliculatus. A, J, K, Torna Fort, Pune Dist., Maharashtra (BMNH 20120038). B, C. conicus W.T. Blanford, 1870, lectotype, Torna Fort, Pune Dist., Maharashtra (BMNH 1906.1.1.2239). C, Torna Fort, Pune Dist., Maharashtra (BMNH 1906.1.1.2243; from the same lot as probable paralectotypes of C. canaliculatus). D, E, Anculotus carinatus Layard, 1855, lectotype, Mahabaleshwar, Satara Dist., Maharashtra (BMNH 20120031). F–I, Torna Fort, Pune Dist., Maharashtra (USNM 317694). L, M, 6 km west of Mahabaleshwar, Satara Dist., Maharashtra (two views of same specimen; ZSI/WGRS/IR.INV-2295). N, O, 44 km west of Bhor, Pune Dist., Maharashtra (ZSI/WGRS/IR.INV-2296, 2297). P, Q, C. conicus var. canaliculatus W.T. Blanford, 1870, lectotype, Torna Fort, Pune Dist., Maharashtra (BMNH 1906.1.1.2240). R, C. conicus var. canaliculatus W.T. Blanford, 1870, probable paralectotype, Torna Fort, Pune Dist., Maharashtra (BMNH 1906.1.1.2243). S, C. fairbanki ‘[W.T.] Blanford’ Hanley &amp; Theobald, 1876, neotype, no locality (BMNH 1906.1.1.2245). T, Mahabaleshwar, Satara Dist., Maharashtra (BMNH 20120039). U, V, Torna Fort, Pune Dist., Maharashtra (USNM 317694). W–Y, Torna Fort, Pune Dist., Maharashtra (USNM 317695). Z, Mahabaleshwar, Satara Dist., Maharashtra (BMNH 20120040). AA, BB, Dongarwadi, Tamhini, Pune Dist., Maharashtra (two views of same specimen; ZSI/WGRS/ IR.INV-2299). CC, DD, Dongarwadi, Tamhini, Pune Dist., Maharashtra (BMNH 20120041). EE, 10 km west of Mahabaleshwar, Raigad Dist., Maharashtra (ZSI/WGRS/ IR.INV-2300).</p> <p>slightly swollen distally, large basal drum-shaped gland, slender filament. Western Maharashtra State.</p> <p>Shell (Figs 4D–E, 9A–O): Shell H 5.0– 12.1 mm. Shape (Table 1) turbinate to high-turbinate; whorls well rounded, not or only weakly angled at periphery; suture moderately or strongly impressed; apex often eroded. Columella broad. Pseudumbilicus usually absent, or a narrow chink (to 0.2 mm), or a hollowed area up to 1.2 mm wide (never perforated) with sharply angled margin continuous with apertural margin (Fig. 9E). Surface lacks distinct ribs, but a ·</p> <p>narrow flattened area may be present at suture, which is occasionally bounded by a weak rib (Fig. 9B, G, H). Surface with satin sheen; fine weak microstriae, sometimes obsolete (Fig. 4D, E). Diameter of first whorl 0.47 mm (N = 1). Colour: mid-brown, olive brown or golden brown, sometimes with indistinct darker bands, or up to four prominent narrow redbrown bands (at suture, mid whorl, just below periphery and around columella; Fig. 9L, N); columella white or purplish; aperture white to yellow-brown, with purple-brown bands showing through.</p> <p>Animal: Head, tentacles and sides of foot grey to black, snout sometimes paler (Fig. 2H). Gills: 23–43 leaflets; black or unpigmented. Operculum (Table 1; Fig. 5E, F): opercular ratio 0.345 –0.423; weakly to moderately calcified, flexible, concave, internal ridge weak or absent. Penis (Fig. 10A–H): unpigmented; relatively large, base wrinkled, strap-shaped, slightly bulbous distally (sometimes opaque) in ethanol-fixed specimens, with basal opaque cream drum-shaped gland composed of subepithelial lobules; invagination 70–90% of length of base in ethanol-fixed specimens; filament slender, rarely slightly protruding in ethanol-fixed specimens. Pallial oviduct: as for genus.</p> <p>Radula (Fig. 7C, D): Relative radula length 1.13– 2.45. Rachidian: length/width 1.24–1.40; 5 cusps (+1 outer denticle on either side). Lateral: 5 cusps (+ 1 inner denticle). Inner marginal: 5 cusps (+ 1 outer denticle). Outer marginal: 3–5 cusps. Major cusp of each of 5 central teeth elongate leaf-shaped with rounded tip, or elongate triangular and pointed; other cusps pointed.</p> <p>Range (Fig. 8): Western Maharashtra State, between Karli and Mahabaleshwar (90 km linear distance). Records (Supporting Table S1): Maharashtra State: Karlee Hill Fort, Kurkulla, near Pune (locality tentatively identified as Karli; USNM 317693); Torna Fort,</p> <p>near Pune (BMNH 1906.1.1.2239, 2242, 20120038; USNM 317694, 317696); 44 km W Bhor (ZSI/WGRS/ IR.INV-2296, 2297, 2298); 6 km W Mahabaleshwar (ZSI/WGRS/IR.INV-2295); Mahabaleshwar Hills (BMNH 1871.9.23.149).</p> <p>Habitat and ecology: On basalt cliffs with algal mat, or bare, covered by flowing water film; on pebbles in shallow pool at foot of cliff with moving water. Collected from 645 m (44 km west of Bhor) and 1103 m (6 km west of Mahabaleshwar). Blanford (1870: 11) recorded that it was ‘met with abundantly on the steep slopes of Torna one of the old Deccan hill forts... The specimens were taken from rocks by the sides of the small torrents running down the hill side.’ His concept of the species included C. canaliculatus. Torna Fort is at an altitude of 1400 m (Supporting Table S1).</p> <p>Remarks: Like the two other Cremnoconchus species with which it occurs, C. conicus displays shell variation, and this variability and co-occurrence have resulted in taxonomic confusion. Blanford (1870) described, in addition to the typical form, a ‘ var. canaliculatus ’ also from Torna, while considering C. carinatus from Mahabaleshwar to be distinct. He noted: ‘The canaliculate variety serves to connect the typical form with carinatus, as many specimens have the angle at the periphery more marked than in the typical conicus; but specimens of carinatus are of a somewhat different form, with considerably less swollen whorls. Perhaps all three forms should be considered as varieties of one species, for which, however, the name carinatus, which is not very appropriate even for full grown specimens of the Mahableshwar shell, can scarcely be retained with propriety.’ (Blanford, 1870: 11). Here it is shown that C. canaliculatus is a distinct species, while C. carinatus is a synonym of C. conicus. Two specimens of C. conicus are present among the probable syntypes of C. canaliculatus. To add to the confusion, C. syhadrensis is also found at Torna, although apparently unknown to Blanford and not present in his material from the site. For example, four large lots from Torna, each of 20–100 dry shells, are present in the Evezard Collection (USNM) made in the late 19th century. Presumably following Blanford’s descriptions, these were originally sorted and labelled according to the presence of spiral ribs: ‘ C. syhadrensis ’ (eight spiral ribs; contains C. syhadrensis alone; USNM 317683); ‘ C. conicus vars’ (variable ribs; contains C. syhadrensis and C. canaliculatus; USNM 317695); ‘ C. conicus var. canaliculatus ’ (strong rib at suture; contains C. syhadrensis alone; USNM 317696); ‘ C. conicus ’ (no ribs; contains mainly C. conicus with a few C. canaliculatus; USNM 317694). As discussed below, for each of the three species mixed in these lots the development of ribs and angulation of the whorls are variable. Far more significant for identification are the less readily apparent characters of surface microsculpture and form of the columella and umbilicus (Table 3).</p> <p>The type series of C. carinatus consists of five large (H up to 9.2 mm) shells of thick texture, turbinate form, with clear but fine microstriae, weakly marked suture, slight peripheral angle (certainly not carinate; the specific epithet is indeed inappropriate, as noted by Blanford, 1870) and wide (but imperforate) pseudumbilical area (Fig. 9D, E). These shells differ from Blanford’s C. conicus (his ‘typical form’; Blanford, 1870), which has a thinner texture, taller spire, indistinct microstriae, impressed suture, rounded whorls, and minute pseudumbilical chink (Fig. 9B). The respective type localities of Mahabaleshwar and Torna are 35 km apart. No specimens exactly resembling Layard’s shells were discovered during fieldwork at Mahabaleshwar in 2010. The stream flowing through the modern town of Mahabaleshwar to the ‘Chinaman Falls’ (Fig. 2D) is now badly polluted; if this was the type locality, Cremnoconchus no longer occur there. However, it may be that the type locality was another of the many small streams in the area, or in the vicinity of the small settlement of Old Mahabaleshwar. Smaller shells (H up to 6.6 mm) were collected 6 km from Mahabaleshwar. These are of thinner texture, with four dark bands and closed pseudumbilicus (Fig. 9L, M). A collection from 44 km west of Bhor, midway between Torna and Mahabaleshwar (Fig. 9N, O), includes specimens that overlap the range of variation at the other two sites, supporting the proposed synonymy.</p> <p>There is also some variation in the operculum; those of the specimens from near Bhor are thin, flexible, and slightly less tightly coiled (Fig. 5E, F), but those from near Mahabaleshwar are more heavily calcified, appearing more opaque and drying flat (Table 1).</p> <p>It is not known whether there is some segregation by microhabitat where C. syhadrensis, C. canaliculatus and C. conicus occur together, as at Torna. Only C. conicus has been found in shallow pools, while both it and C. canaliculatus occur in flowing water, and C. syhadrensis appears to favour damp rocks and shady overhangs. Although C. conicus and C. canaliculatus both occur in the vicinity of Mahabaleshwar, in apparently similar habitats, they were not discovered in the same streams (see Supporting Table S1 for locality details).</p> <p>Table 3 summarizes the diagnostic characters of the three Cremnoconchus of western Maharashtra. Cremnoconchus conicus can be identified by its satin sheen, wide columella and thin (mainly corneous) operculum. The penial shape of C. syhadrensis is diagnostic, but those of the other two are more similar. Compared with that of C. canaliculatus the penis of C. conicus is relatively larger, has a more swollen apex to the base, and the drum-shaped glandular projection is larger (Fig. 10).</p> <p>CREMNOCONCHUS CANALICULATUS W.T.</p> </div>	https://treatment.plazi.org/id/038A87FB192301779BD6FEF5E67BFC56	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Reid, David G.;Aravind, Neelavara Ananthram;Madhyastha, Neelavara Ananthram	Reid, David G., Aravind, Neelavara Ananthram, Madhyastha, Neelavara Ananthram (2013): A unique radiation of marine littorinid snails in the freshwater streams of the Western Ghats of India: the genus Cremnoconchus W. T. Blanford, 1869 (Gastropoda: Littorinidae). Zoological Journal of the Linnean Society 167 (1): 93-135, DOI: 10.1111/j.1096-3642.2012.00875.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2012.00875.x
038A87FB1930017C9BFEFE4DE204FD01.text	038A87FB1930017C9BFEFE4DE204FD01.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Cremnoconchus hanumani Reid & Aravind & Madhyastha 2013	<div><p>CREMNOCONCHUS HANUMANI SP. NOV.</p> <p>(FIGS 4N, O, 5I, J, 7G, H, 8, 11A–G, 12A–G)</p> <p>Types: Holotype ZSI / WGRS /IR.INV-2303 (Fig. 11A, B); 1 paratype ZSI / WGRS /IR.INV-2304 (Fig. 11C); <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=75.15511&amp;materialsCitation.latitude=13.27008" title="Search Plazi for locations around (long 75.15511/lat 13.27008)">Hanuman Gundi Falls</a>, Chikmagalur Dist., Karnataka, India (13.27008°N 75.15511°E).</p> <p>Etymology: After the Hindu deity Hanuman, from the type locality.</p> <p>Diagnosis: Shell turbinate to globular, without ribs; pseudumbilicus broad but not perforated; surface with satin sheen, faint microstriae. Operculum weakly calcified, no internal ridge. Penis with slight distal swelling, stout filament. Western Karnataka State.</p> <p>Shell (Figs 4N, O, 11A–G): Shell H 3.3–5.0 mm. Shape (Table 1) globular to turbinate; whorls rounded, not or only slightly angled at periphery; suture impressed, with slightly flattened ramp and weakly angled shoulder; apex eroded; base slightly swollen. Delicate texture. Columella narrow. Pseudumbilicus broad (to 1.0 mm; rarely 0.3–0.5 mm), hollowed but not deeply perforated, outlined by sharply angled margin continuous with apertural margin (Fig. 11B). Surface without ribs; with satin sheen; very fine or faint spiral striae (Fig. 4N, O), becoming obsolete on last whorl of largest shells. Protoconch 1.4 whorls; diameter 0.62 (N = 1); diameter of first whorl 0.45– 0.53 mm (N = 2). Colour: dark brown or olive-brown,</p> <p>› Figure 11. Shells of Cremnoconchus species. A –G, C. hanumani. H–L, C. globulus. M–P, C. agumbensis. Q–U, C. cingulatus. V–AA, C. castanea. BB–EE, C. dwarakii. A–C, C. hanumani sp. nov., holotype (A, B; ZSI/WGRS/IR.INV-2303) and paratype (C; ZSI/WGRS/IR.INV-2304), Hanuman Gundi Falls, Chikmagalur Dist., Karnataka. D–G, Greater Kadambi Falls, Chikmagalur Dist., Karnataka (D, E two views of same specimen; ZSI/WGRS/IR.INV-2305, 2306, 2307). H, L, Greater Kadambi Falls, Chikmagalur Dist., Karnataka (ZSI/WGRS/IR.INV-2308, 2309). I–K, C. globulus sp. nov., holotype (I, J; ZSI/WGRS/IR.INV-2310) and paratype (K; ZSI/WGRS/IR.INV-2311), Lesser Kadambi Falls, Chikmagalur Dist., Karnataka. M–P, C. agumbensis sp. nov., holotype (M, N; ZSI/WGRS/IR.INV-2313) and paratypes (O, P; ZSI/WGRS/IR.INV-2314, 2315), Someshwara to Agumbe road, Udupi Dist., Karnataka. Q–U, C. cingulatus sp. nov., holotype (R, S; ZSI/WGRS/IR.INV-2316) and paratypes (Q, T, U; ZSI/WGRS/IR.INV-2317, 2318, 2319), Hulikal Ghat, Udupi Dist., Karnataka. V–X, C. castanea sp. nov., holotype (W, X; ZSI/WGRS/IR.INV-2321) and paratype (V; ZSI/ WGRS/IR.INV-2320), Belkal Thirtha Falls, Udupi Dist., Karnataka. Y, Arasinagundi Falls, Udupi Dist., Karnataka (BMNH 20120032). Z, AA, no locality (two views of same specimen; ZSI/WGRS/IR.INV-2322). BB–DD, C. dwarakii sp. nov., holotype (BB, CC; ZSI/WGRS/IR.INV-2323) and paratype (DD; ZSI/WGRS/IR.INV-2324), Hulikal Ghat, Udupi Dist., Karnataka. EE, Hulikal Ghat, Udupi Dist., Karnataka (BMNH 20120036).</p> <p>sometimes a narrow purple-brown band at suture; columella whitish or slightly tinged purple-brown; aperture pale with sutural band showing through.</p> <p>Animal: Head, tentacles, and sides of foot dark grey to black. Gills: 24–30 leaflets; grey or unpigmented. Operculum (Table 1; Fig. 5I, J): opercular ratio 0.318 – 0.400; weakly calcified, translucent mid-brown, no internal ridge. Penis (Fig. 12A–G): unpigmented or slightly pigmented; base wrinkled, slightly swollen (possibly glandular) distally; invagination 70–90% of length of base in ethanol-fixed specimens; filament relatively stout, usually protruding in ethanol-fixed specimens. Pallial oviduct: as for genus. One pallial oviduct contained single egg with firm covering 0.47 mm diameter (ethanol fixed).</p> <p>Radula (Fig. 7G, H): Relative radula length 2.50– 3.06. Rachidian: length/width 1.09–1.21; 5 cusps (+ 1 outer denticle on either side). Lateral: 5 cusps (+ 1–2 inner denticles). Inner marginal: 6 cusps. Outer marginal: 6–7 cusps (+ 1 inner or outer denticle). Major cusp of each of 5 central teeth leaf-shaped with pointed tip; other cusps pointed.</p> <p>Range (Fig. 8): Western Karnataka State, Kudremukh (55 km north-east of Mangalore). Records (Supporting Table S1): Karnataka State: Hanuman Gundi Falls (ZSI/WGRS/IR.INV-2303, 2304); Greater Kadambi Falls (ZSI/WGRS/IR.INV-2305, 2306, 2307).</p> <p>Habitat and ecology: On rocks and cliff wetted by spray from strong waterfall; in crevices; in partial</p> <p>shade of riparian vegetation in wet evergreen forest. Altitude 830 m and 941 m.</p> <p>Remarks: The small, globose shell with delicate texture and wide (but not perforated) pseudumbilicus are distinctive of this species. Cremnoconchus castanea has a taller, thicker shell and well-calcified operculum (Table 4). Cremnoconchus globulus can be found together with C. hanumani (Supporting Table S1); the former is larger, more solid, with a moderate umbilicus; the operculum is weakly calcified in both, but dark red-brown in C. globulus and translucent mid-brown in C. hanumani. The penis of C. hanumani is diagnostic.</p> <p>Where this species occurs in the same stream as the larger C. globulus there is no obvious difference in microhabitat.</p> </div>	https://treatment.plazi.org/id/038A87FB1930017C9BFEFE4DE204FD01	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Reid, David G.;Aravind, Neelavara Ananthram;Madhyastha, Neelavara Ananthram	Reid, David G., Aravind, Neelavara Ananthram, Madhyastha, Neelavara Ananthram (2013): A unique radiation of marine littorinid snails in the freshwater streams of the Western Ghats of India: the genus Cremnoconchus W. T. Blanford, 1869 (Gastropoda: Littorinidae). Zoological Journal of the Linnean Society 167 (1): 93-135, DOI: 10.1111/j.1096-3642.2012.00875.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2012.00875.x
038A87FB1935017C98A8FD59E6A3FABA.text	038A87FB1935017C98A8FD59E6A3FABA.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Cremnoconchus globulus Reid & Aravind & Madhyastha 2013	<div><p>CREMNOCONCHUS GLOBULUS SP. NOV.</p> <p>(FIGS 2E, 4F, 5K, L, 8, 11H–L, 12H–L, 13A, B)</p> <p>Types: Holotype ZSI / WGRS /IR.INV-2310 (Fig. 11I, J); 2 paratypes ZSI / WGRS /IR.INV-2311, 2312 (Fig. 11K); <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=75.17056&amp;materialsCitation.latitude=13.24384" title="Search Plazi for locations around (long 75.17056/lat 13.24384)">Lesser Kadambi Falls</a>, Chikmagalur Dist., Karnataka, India (13.24384°N, 75.17056°E).</p> <p>Etymology: Latin globulus, globular, in reference to shell shape.</p> <p>Diagnosis: Shell globular, without ribs; pseudumbilicus moderate, sometimes perforated; surface with satin sheen, no microstriae. Operculum weakly calcified, no internal ridge. Penis with lateral glandular flange, slender filament. Western Karnataka State.</p> <p>Shell (Figs 4F, 11H–L): Shell H 6.0– 8.8 mm. Shape (Table 1) globular; whorls rounded, without angulation; suture impressed; apex eroded; base slightly swollen. Columella moderately narrow, wider at base. Pseudumbilicus moderate (to 0.8 mm), sometimes perforated, outlined by angled margin, sometimes forming a slight rounded rib. Surface almost always without ribs above periphery; rarely a slight thickening or indistinct rib near suture. Surface with satin sheen; spiral striae almost or entirely absent (Fig. 4F). Diameter of first whorl 0.50–0.66 mm (N = 3). Colour: dark brown or olive-brown, sometimes darker on spire and in band at suture; columella and umbilicus purple-brown; aperture pale brown to whitish, with sutural band showing through.</p> <p>Animal: Head, tentacles, and sides of foot pale grey to black, tentacles darker, paler at tip of snout. Gills: up to 40 leaflets; black. Operculum (Table 1; Fig. 5K, L): opercular ratio 0.364 –0.421; weakly calcified, dark red-brown, no internal ridge. Penis (Fig. 12H–L): unpigmented or slightly pigmented; base wrinkled, with long thickened flange running across left side towards eye, glandular knob on right side, and slight glandular swelling distally (sometimes opaque); invagination about half length of base in ethanol-fixed specimens; filament slender, rarely protruding in ethanol-fixed specimens. Pallial oviduct: as for genus.</p> <p>Radula (Fig. 13A, B): Relative radula length 2.66– 3.43. Rachidian: length/width 1.10–1.22; 5 cusps (+ 1 outer denticle on either side). Lateral: 5 cusps (+ 1 inner denticle). Inner marginal: 5 cusps. Outer marginal: 4–5 cusps. Major cusp of each of 5 central teeth triangular leaf-shaped with pointed to slightly rounded tip; other cusps pointed.</p> <p>Range (Fig. 8): Western Karnataka State, Kudremukh (55 km north-east of Mangalore). Records (see Supporting Table S1): Karnataka State: Lesser Kadambi Falls (ZSI/WGRS/IR.INV-2310, 2311, 2312); Greater Kadambi Falls (ZSI/WGRS/IR.INV-2308, 2309).</p> <p>Habitat and ecology: Common in film of water flowing over rock face beside waterfall (Fig. 2E); on stones in shallow streams (to 30 cm deep) with fast-flowing water; in partial shade of riparian vegetation in wet evergreen forest. Altitude 941 and 967 m.</p> <p>Remarks: Four Cremnoconchus species have similar umbilicate, globular to turbinate, smooth shells (Table 4). Cremnoconchus globulus is distinguished by its lack of a basal rib (present in C. cingulatus), moderate pseudumbilicus and weakly calcified operculum (well calcified in C. castanea). Distinction from C. hanumani, with which it occurs in the same microhabitat, is discussed in the Remarks on that species. The penes of all four are diagnostic (Figs 12, 14, 15).</p> </div>	https://treatment.plazi.org/id/038A87FB1935017C98A8FD59E6A3FABA	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Reid, David G.;Aravind, Neelavara Ananthram;Madhyastha, Neelavara Ananthram	Reid, David G., Aravind, Neelavara Ananthram, Madhyastha, Neelavara Ananthram (2013): A unique radiation of marine littorinid snails in the freshwater streams of the Western Ghats of India: the genus Cremnoconchus W. T. Blanford, 1869 (Gastropoda: Littorinidae). Zoological Journal of the Linnean Society 167 (1): 93-135, DOI: 10.1111/j.1096-3642.2012.00875.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2012.00875.x
038A87FB1935017F9B1EFA12E0E2FEB3.text	038A87FB1935017F9B1EFA12E0E2FEB3.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Cremnoconchus agumbensis Reid & Aravind & Madhyastha 2013	<div><p>CREMNOCONCHUS AGUMBENSIS SP. NOV.</p> <p>(FIGS 4H, 5M, N, 8, 11M–P, 13C, D, 14A–C, 16E)</p> <p>Types: Holotype ZSI / WGRS /IR.INV-2313 (Fig. 11M, N); 2 paratypes ZSI / WGRS /IR.INV-2314, 2315 (Fig. 11O, P); <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=75.07783&amp;materialsCitation.latitude=13.49342" title="Search Plazi for locations around (long 75.07783/lat 13.49342)">Someshwara</a> to Agumbe road, Udupi Dist., Karnataka, India (13.49342°N, 75.07783°E).</p> <p>Etymology: From the type locality.</p> <p>Diagnosis: Shell high turbinate, shoulder angled or with rib; pseudumbilicus absent; surface dull, with strong microstriae. Operculum calcified, no internal ridge. Penis large, with distal glandular thickening, stout filament. Western Karnataka State.</p> <p>Shell (Figs 4H, 11M–P): Shell H 5.5–10.4 mm. Shape (Table 1) high turbinate; whorls strongly angled at shoulder (sometimes raised as a rib; Fig. 11N), giving pronounced sutural ramp or channel, but becoming weak at end of last whorl; slight angle at periphery; suture impressed; apex slightly eroded; base slightly swollen. Columella narrow. Pseudumbilicus absent. Surface with single narrow rib at shoulder; dull, with irregular but distinct microstriae, fainter on last whorl and on base (Fig. 4H). Protoconch (Fig. 16E) 1.5 whorls; diameter 0.68 mm; diameter of first whorl 0.53 mm (n = 1). Colour: dark brown, darker on spire, three more or less distinct broad black-brown bands at shoulder, above periphery and on inner part of base; aperture pale brown, columella and interior bands purple-brown.</p> <p>Animal: Head, tentacles and sides of foot black, paler at tip of snout. Gills: up to 50 leaflets; unpigmented. Operculum (Table 1; Fig. 5M, N): opercular ratio 0.439 –0.565; calcified, no internal ridge. Penis (Fig. 14A, B): relatively large; unpigmented; base wrinkled, strap-shaped, with slightly thickened distal glandular region; filament relatively stout, protruding in ethanol-fixed specimens. Pallial oviduct (Fig. 14C): as for genus. One pallial oviduct contained two eggs 0.40 mm diameter (ethanol fixed).</p> <p>Radula (Fig. 13C, D): Relative radula length 1.09– 1.79. Rachidian: length/width 1.19–1.42; 5 cusps (+ 1 outer denticle on either side). Lateral: 6 cusps (sometimes + 1 inner denticle). Inner marginal: 5 cusps (sometimes + 1 inner denticle). Outer marginal: 4 cusps (sometimes + 1 inner denticle). Major cusp of each of 5 central teeth elongate-triangular with rounded and slightly papillose tip; other cusps pointed.</p> <p>Range (Fig. 8): Western Karnataka State, Agumbe (75 km north-northeast of Mangalore). Records (Supporting Table S1): Karnataka State: Someshwara to Agumbe road (ZSI / WGRS /IR.INV-2313, 2314, 2315).</p> <p>Habitat and ecology: Damp rock face shaded by evergreen forest. Altitude 379 m.</p> <p>Remarks: Among the known Cremnoconchus species of Karnataka, C. agumbensis is the only one with turreted whorls produced by a sharp angulation (or rib) at the shoulder, and without a pseudumbilicus (Table 4). In these characters it resembles the canaliculate form of C. canaliculatus from Maharashtra (Table 3; Fig. 9P, R, U–Z, CC), but that has a more solid shell and the penis bears a drum-shaped gland (Fig. 10I–N).</p> </div>	https://treatment.plazi.org/id/038A87FB1935017F9B1EFA12E0E2FEB3	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Reid, David G.;Aravind, Neelavara Ananthram;Madhyastha, Neelavara Ananthram	Reid, David G., Aravind, Neelavara Ananthram, Madhyastha, Neelavara Ananthram (2013): A unique radiation of marine littorinid snails in the freshwater streams of the Western Ghats of India: the genus Cremnoconchus W. T. Blanford, 1869 (Gastropoda: Littorinidae). Zoological Journal of the Linnean Society 167 (1): 93-135, DOI: 10.1111/j.1096-3642.2012.00875.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2012.00875.x
038A87FB193601419B09FE17E122FB2A.text	038A87FB193601419B09FE17E122FB2A.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Cremnoconchus cingulatus Reid & Aravind & Madhyastha 2013	<div><p>CREMNOCONCHUS CINGULATUS SP. NOV.</p> <p>(FIGS 4G, 5O, P, 11Q–U, 13E, F, 14D–I)</p> <p>Types: Holotype ZSI / WGRS /IR.INV-2316 (Fig. 11 R, S); 3 paratypes ZSI / WGRS /IR.INV-2317, 2318, 2319 (Fig. 11Q, T, U); <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=74.99445&amp;materialsCitation.latitude=13.71742" title="Search Plazi for locations around (long 74.99445/lat 13.71742)">Hulikal Ghat</a>, Udupi Dist., Karnataka, India (13.71742°N 74.99445°E).</p> <p>Etymology: Latin cingulatus, girdled, in reference to sculpture of base.</p> <p>Diagnosis: Shell globular; one rib on base; pseudumbilicus narrow, perforated; surface with satin sheen, microstriae weak or absent. Operculum weakly calcified, no internal ridge. Penis with distal glandular pad, slender filament. Western Karnataka State.</p> <p>Shell (Figs 4G, 11Q–U): Shell H 3.5–6.7 mm. Shape (Table 1) globular; whorls rounded; a weak rib close to suture outlines the flattened sutural area, but rib becomes obsolete on largest shells; suture impressed; peripheral angle absent; apex eroded; base slightly swollen, with thick rib at one-third radius. Columella narrow, wider at base. Pseudumbilicus narrow, 0.2– 0.6 mm, perforated. Surface with satin sheen; spiral striae indistinct or absent, weakly present on base (Fig. 4G). Colour: reddish brown, three indistinct darker brown bands at suture, above periphery and at middle of base; aperture and columella purple-brown, with bands showing through.</p> <p>Animal: Head and tentacles dark grey to black, tip of snout paler; body and sides of foot black. Gills: up to 35 leaflets; black. Operculum (Table 1; Fig. 5O, P): opercular ratio 0.334 –0.472; weakly calcified, concave, no internal ridge. Penis (Fig. 14D–I): base lightly pigmented, wrinkled, solid, with large glandular pad on left of reverse side; invagination about half length of base in ethanol-fixed specimens; filament slender, protruding in ethanol-fixed specimens. Pallial oviduct: as for genus.</p> <p>› Figure 14. Anatomy of Cremnoconchus agumbensis (A–C) and C. cingulatus (D–I), all to same magnification. A, B, D–I, penes (all fixed in ethanol); G–I are abaxial, sectioned and adaxial views of same penis. C, pallial oviduct, with transverse sections in three positions. A–C, Agumbe, Udupi Dist., Karnataka (ZSI/WGRS; shell H: A = 9.5 mm; B = 8.8 mm; C = 9.1 mm). D–I, Hulikal Ghat, Udupi Dist., Karnataka (ZSI/WGRS; shell H: D = 3.5 mm; E = 4.6 mm; F = 3.7 mm; G–I = 4.4 mm). Shading conventions as in Figure 6.</p> <p>Radula (Fig. 13E, F): Relative radula length 1.93– 2.09. Rachidian: length/width 1.10–1.13; 5 cusps (+ 1 outer denticle on either side). Lateral: 5 cusps (sometimes + 1 inner denticle). Inner marginal: 5 cusps. Outer marginal: 4–5 cusps. Major cusp of each of 5 central teeth triangular with rounded and slightly papillose tip; other cusps pointed.</p> <p>Range (Fig. 8): Western Karnataka State, Hulikal Ghat (95 km north-northeast of Mangalore). Records (Supporting Table S1): Karnataka State: Hulikal Ghat, Udupi District (ZSI / WGRS /IR.INV-2316, 2317, 2318, 2319).</p> <p>Habitat and ecology: In rushing stream on rock face and sides of boulders, in fast-flowing water and spray, partly shaded by trees (Fig. 2C). Altitude 475 m.</p> <p>Remarks: The conspicuous rib on the base, narrow but deep pseudumbilicus, and concave, weakly calcified operculum are the distinguishing features of the shell of this species. Except for the basal rib, these characters resemble C. globulus, but their respective penes are diagnostic (Figs 12H–L, 14D–I). Characters of the Cremnoconchus species of Karnataka are summarized in Table 4.</p> <p>There appears to be sexual dimorphism: shells of males are smaller (males: mean ± sample SD = 3.840 ± 0.619 mm, N = 4; females: 5.575 ± 0.697 mm, N = 12) and the aperture is slightly enlarged.</p> <p>At Hulikal Ghat it was found together with C. dwarakii and the two occupied different microhabitats. Cremnoconchus cingulatus occurred on rocks in rushing water, while C. dwarakii was found on mossy rocks dampened only by spray.</p> </div>	https://treatment.plazi.org/id/038A87FB193601419B09FE17E122FB2A	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Reid, David G.;Aravind, Neelavara Ananthram;Madhyastha, Neelavara Ananthram	Reid, David G., Aravind, Neelavara Ananthram, Madhyastha, Neelavara Ananthram (2013): A unique radiation of marine littorinid snails in the freshwater streams of the Western Ghats of India: the genus Cremnoconchus W. T. Blanford, 1869 (Gastropoda: Littorinidae). Zoological Journal of the Linnean Society 167 (1): 93-135, DOI: 10.1111/j.1096-3642.2012.00875.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2012.00875.x
038A87FB190801439882FA84E239FCE7.text	038A87FB190801439882FA84E239FCE7.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Cremnoconchus castanea Reid & Aravind & Madhyastha 2013	<div><p>CREMNOCONCHUS CASTANEA SP. NOV.</p> <p>(FIGS 4M, 5Q, R, 11V–AA, 13G, H, 15A–D)</p> <p>Types: Holotype ZSI / WGRS /IR.INV-2321 (Fig. 11W, X); 1 paratype ZSI / WGRS /IR.INV-2320 (Fig. 11 V); <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=74.897&amp;materialsCitation.latitude=13.842" title="Search Plazi for locations around (long 74.897/lat 13.842)">Belkal Thirtha Falls</a>, near Kollur, Udupi Dist., Karnataka, India (13.842°N 74.897°E).</p> <p>Etymology: Latin castanea, a chestnut, in reference to shell colour, used as a noun in apposition.</p> <p>Diagnosis: Shell turbinate; ribs absent; pseudumbilicus wide, perforated; surface with indistinct or strong microstriae. Operculum calcified, with internal ridge. Penis large, with distal glandular pad, slender filament. Western Karnataka State.</p> <p>Shell (Figs 4M; 11V–AA): Shell H 4.7–9.4 mm. Shape turbinate; whorls slightly angled at shoulder and periphery; ribs absent; suture impressed; apex eroded; base slightly swollen. Columella narrow. Pseudumbilicus broad, 0.5–1.0 mm, with sharply angled margin, perforated. Surface dull or with satin sheen; indistinct or strong, coarse microstriae cover surface (Fig. 4M). Diameter of first whorl 0.32– 0.53 mm (N = 2). Colour: orange-brown to chestnut brown, darker on spire and with indistinct darker band at suture; aperture pale brown, columella pinkish-brown to white.</p> <p>Animal: Head, tentacles, sides of foot, and body pale grey to black. Gills: unpigmented. Operculum (Table 1; Fig. 5Q–R): opercular ratio 0.341 –0.450; well calcified, flat, with internal ridge. Penis (Fig. 15A–D): relatively large, unpigmented; base wrinkled, elongate, solid, with large triangular glandular pad on reverse side; invagination about half length of base in ethanol-fixed specimens; filament slender, protruding in ethanolfixed specimens. Pallial oviduct: as for genus.</p> <p>Radula (Fig. 13G, H): Relative radula length 2.37– 2.59. Rachidian: length/width 1.08–1.26 (–1.64); 5 cusps (+ 1 outer denticle on either side). Lateral: 5 cusps (+ 1 inner denticle). Inner marginal: 5 cusps (+ 1 inner denticle). Outer marginal: 4–5 cusps. Major cusp of each of 5 central teeth leaf-shaped with pointed or slightly papillose tip; other cusps pointed.</p> <p>Range (Fig. 8): Western Karnataka State, near Kollur (110 km north of Mangalore). Records (Supporting Table S1): Karnataka State: Belkal Thirtha Falls (ZSI/WGRS/IR.INV-2320, 2321); Arasinagundi Falls (BMNH 20120032).</p> <p>Habitat and ecology: Spray zone of strong waterfalls. Large groups found aestivating in crevices during summer at Arasinagundi Falls. Altitude 373– 384 m.</p> <p>Remarks: Four species (C. castanea, C. cingulatus, C. globulus, C. hanumani) from Karnataka show similarities in their turbinate to globular shells</p> <p>› Figure 15. Anatomy of Cremnoconchus castanea (A–D) and C. dwarakii (E–J), all to same magnification. A–D, E–I, penes (A, live, relaxed; B, C, live, contracted; D–I, fixed in ethanol); B, C, are abaxial and adaxial views of same penis; G, H, are abaxial and sectioned views of same penis. J, pallial oviduct. A–C, Arasinagundi Falls, Udupi Dist., Karnataka (BMNH 20120034; shell H: A = 7.6 mm; B, C = 4.7 mm). D, Belkal Thirtha Falls, Udupi Dist., Karnataka (ZSI/WGRS; shell H = 7.0 mm). E–J, Hulikal Ghat, Udupi Dist., Karnataka (ZSI/WGRS; shell H: E = 6.4 mm; F = 6.2 mm; G, H = 6.1 mm; I = 6.9 mm; J = 6.9 mm). Shading conventions as in Figure 6.</p> <p>without ribs on their upper surface. Details of the relative size of the pseudumbilicus and degree of calcification of the operculum can aid discrimination, but penial shape is most useful (Table 4). Of these four, only C. castanea is known to develop strong, coarse microstriae, but these can sometimes be indistinct. The combination of large umbilicus and strong microstriae is similar to some examples of C. canaliculatus from Maharashtra (Fig. 9AA–EE), but that species has a basal drum-shaped gland on the penis (Fig. 10I–N).</p> <p>An additional unlocalized sample (ZSI/WGRS/ IR.INV-2322) could belong to this species. The shells (Fig. 11Z, AA) differ in that the pseudumbilicus, while perforated, is narrower than in the shells described above (0.3–0.5 mm in shells with H = 4.3–5.6 mm; Fig. 11V–Y) and has a rounded (not sharp) edge. Shell and operculum are otherwise similar. Only a single penis was seen and this was similar to those described.</p></div> 	https://treatment.plazi.org/id/038A87FB190801439882FA84E239FCE7	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Reid, David G.;Aravind, Neelavara Ananthram;Madhyastha, Neelavara Ananthram	Reid, David G., Aravind, Neelavara Ananthram, Madhyastha, Neelavara Ananthram (2013): A unique radiation of marine littorinid snails in the freshwater streams of the Western Ghats of India: the genus Cremnoconchus W. T. Blanford, 1869 (Gastropoda: Littorinidae). Zoological Journal of the Linnean Society 167 (1): 93-135, DOI: 10.1111/j.1096-3642.2012.00875.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2012.00875.x
038A87FB190A0144988FFCF2E17AF8A9.text	038A87FB190A0144988FFCF2E17AF8A9.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Cremnoconchus dwarakii Reid & Aravind & Madhyastha 2013	<div><p>CREMNOCONCHUS DWARAKII SP. NOV.</p> <p>(FIGS 4A, 5C, D, 11BB–EE, 15E–J, 16A, B, F)</p> <p>Cremnoconchus syhadrensis – Williams et al., 2003: 63 (not Blanford, 1863).</p> <p>Cremnoconchus dwaraki Madhyastha, 2008: The Hindu 29 June 2008 (as Cremenoconchus; nomen nudum).</p> <p>Taxonomic history: This species was misidentified by Williams et al. (2003) as C. syhadrensis, because the two share strong spiral ribs on the shell.</p> <p>Types: Holotype ZSI / WGRS /IR.INV-2323 (Fig. 11 BB, CC); four paratypes ZSI / WGRS /IR.INV-2324, 2325 (Fig. 11DD); <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=74.99445&amp;materialsCitation.latitude=13.71742" title="Search Plazi for locations around (long 74.99445/lat 13.71742)">Hulikal Ghat</a>, Udupi District, Karnataka, India (13.71742°N, 74.99445°E).</p> <p>Etymology: Named after Acharya Dwarakanth, of the Indian Naturalist Club. He was known for his knowledge of animals and plants of Karnataka and, as secretary of the club, supported many research activities and encouraged the younger generation in birding and nature conservation. The name dwarakii is derived from that of the legendary city of Dwaraka in Western Gujarat, the dwelling place of Lord Krishna and the capital of the Yadus who ruled the Anarta Kingdom during the Mahabaratha period.</p> <p>Diagnosis: Shell turbinate; 8–11 ribs (including base); pseudumbilicus narrow, perforated; surface dull, with fine strong microstriae. Operculum weakly calcified, no internal ridge. Penis with slight glandular flange, slender filament. Western Karnataka State. COI: GenBank AJ 488605.</p> <p>Shell (Figs 4A, 11BB–EE): Shell H 6.0– 9.2 mm. Shape (Table 1) turbinate; whorls slightly angled at shoul- der; suture impressed; apex slightly eroded; base slightly swollen. Columella narrow. Pseudumbilicus narrow, 0.2–0.7 mm, with sharply angled margin, perforated. Spire whorls with three strong ribs. Last whorl with 8–11 strong ribs, of which largest are sutural rib (outlining flattened sutural ramp, which may also bear an additional riblet) and rib close to pseudumbilicus. Surface dull, covered by fine, strong microstriae, intersecting with fine growth lines to give minutely lamellose effect (Fig. 4A). Protoconch (Fig. 16F) 1.4 whorls; diameter 0.70 mm; diameter of first whorl 0.45 mm (N = 1). Colour: usually orangebrown, darker brown on spire and with indistinct darker band at suture and outlining pseudumbilicus; occasionally all rib interspaces are dark brown, but fading on last whorl; aperture yellow-brown, columella and pseudumbilicus purple-brown, exterior bands showing through.</p> <p>Animal: Head, tentacles, sides of foot, and body dark grey to black; tip of snout pale. Gills: up to 36 leaflets; grey to black. Operculum (Table 1; Fig. 5C, D): opercular ratio 0.385 –0.511; weakly calcified, no internal ridge, slightly concave. Penis (Fig. 15E–I): unpigmented; base wrinkled, tapering to rounded tip; with triangular glandular pad on reverse side, slightly projecting anterolaterally; invagination about half length of base in ethanol-fixed specimens; filament slender, not protruding in ethanol-fixed specimens. Pallial oviduct (Fig. 15J): as for genus. One pallial oviduct contained two eggs 0.40 mm in diameter (ethanol fixed), one at two-cell embryo stage.</p> <p>Radula (Fig. 16A, B): Relative radula length 0.82– 1.25. Rachidian: length/width 1.23; 5 cusps (+ 1 outer denticle on either side). Lateral: 5 cusps (+ 1 inner denticle). Inner marginal: 6 cusps. Outer marginal: 3–5 cusps. Major cusp of each of 5 central teeth leaf-shaped with narrowly rounded or slightly papillose tip; other cusps pointed.</p> <p>Range (Fig. 8): Western Karnataka State, Hulikal Ghat (95 km north-northeast of Mangalore). Records (Supporting Table S1): Karnataka State: Hulikal Ghat, Udupi District (ZSI/WGRS/IR.INV-2323, 2324, 2325; BMNH 20120036).</p> <p>Habitat and ecology: On mossy rocks dampened by spray from rushing stream; shaded by trees (Fig. 2B). Altitude 475 m.</p> <p>Remarks: This species is distinguished from all others in Karnataka by the strong ribs of the shell (Table 4). These create a superficial resemblance to ribbed forms of C. syhadrensis from Maharashtra (Fig. 3), but in those the ribs are restricted to the upper surface (i.e. above the periphery), the pseudumbilicus is usually wide and the columella is broad; there are also differences in the penes of these two species (Table 3, Fig. 6A, B, D–G, 15E–I).</p> <p>At Hulikal Ghat this species was found with C. cingulatus (see Remarks on that species).</p> </div>	https://treatment.plazi.org/id/038A87FB190A0144988FFCF2E17AF8A9	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Reid, David G.;Aravind, Neelavara Ananthram;Madhyastha, Neelavara Ananthram	Reid, David G., Aravind, Neelavara Ananthram, Madhyastha, Neelavara Ananthram (2013): A unique radiation of marine littorinid snails in the freshwater streams of the Western Ghats of India: the genus Cremnoconchus W. T. Blanford, 1869 (Gastropoda: Littorinidae). Zoological Journal of the Linnean Society 167 (1): 93-135, DOI: 10.1111/j.1096-3642.2012.00875.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2012.00875.x
