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We have developed a strategy for selective imaging of collagen in live foetal ovine osteoblasts. Our approach involves the incorporation of an azide-tagged proline in the biosynthesis of collagen followed by labelling using a strain-promoted [3+2] azide-alkyne cycloaddition reaction.

The 2019 meeting of WGECO was held at the International Council for the Exploration of the Sea, Copenhagen, Denmark from 8–16 April 2019. The meeting was attended by 16 delegates from eight countries and was chaired by Stefán Ragnarsson (Iceland) and Jeremy Collie (USA). The work conducted centred on three Terms of Reference that had been made by WGECO. In addition, a list of sensitive species prepared by WGBYC was reviewed. WGECO continued the work initiated in 2018 to examine the ecological consequences of stock rebuilding, with emphasis on benthivorous fish (ToRa). Two case studies were carried out to estimate the predation pressure of fish on benthos. In one, the consumption of benthic invertebrates by demersal fish on the US continental shelf regions was compared with their biomass in sediments. The estimated annual consumption of benthic organisms was a small proportion (WGECO scrutinized methods to estimate density-dependent effects on fish growth and made suggestions on its estimation from stock assessment or survey data. A simulation study showed that the age-specific growth increment may not be a useful variable to test for density-dependent growth, because it depends on body size. In the WGECO 2018 report, the case had been made that North Sea plaice show density-dependent growth reduction. In 2019, we conducted a more thorough analysis of the spatial and temporal trends in distribution, growth and status of plaice in the North Sea. As the plaice population increased in abundance, the spatial distribution did not expand, apart from an increased occupancy of marginal areas. The observed reduction in plaice weight-at-length may thus be explained by the metabolic stress associated with the effects of rising sea temperature on plaice growth rates. However, the two mechanisms, temperature and density, that can explain reduced weight-at-age are not mutually exclusive. As part of ToR b, WGECO compiled fisheries-dependent size composition data from four regions: West of Scotland, Irish Sea, West of Ireland, and the Northern Celtic Sea. This novel dataset was used to examine changes in the length composition of the catch in relation to changes in total catch (used as a proxy for fishing pressure). In the Irish Sea, for example, the commercial fishery now catches larger individuals of larger demersal species as the total catch has declined. Size composition of the landings was compared with size composition in the surveys. In the northern Celtic Sea and West of Scotland, the commercial fleets generally take fish of a much larger size than those caught by the survey. By contrast, in the Irish Sea the commercial fleets tend to capture fish across the whole size range available in the environment. Demersal fish community indicators of species composition and size–structure were investigated for several sea areas, using survey data for state, and fisheries-dependent data for pressure. Together, these indicators can be used to track the wider impacts of fisheries on the ecosystem and monitor the evolving nature of the relationship between pressure and state. Simple empirical approaches for setting targets and baselines for demersal fish communities were tested with sensitivity analyses and applied to additional survey areas. WGECO recommends that these indicators be adopted by ICES for evaluating acceptable status for communities and supplemented where possible with risk-based targets from multispecies modelling approaches. A multispecies model framework was advanced to determine more ambitious risk-based targets for demersal fish community indicators that can be used to minimise the risk that stocks suffer from impaired reproductive capacity consistent with commercial fish stock indicators. Ensemble modelling demonstrated that the fish community in the North Sea is recovering in response to fisheries management. WGECO reviewed potential indicators of spatial distribution with special attention to the simplicity of use and ease of communicating the results to non-experts (ToRc). Metrics including aspects of latitude, longitude or depth (e.g. range and centre of gravity) and the area containing a fixed percentage of the population (termed core area) were considered simple to use and easy to communicate. Other measures were considered useful for scientific purposes but less easily applied and communicated to managers. The accuracy of selected indicators for spatial distribution were explored in a simulation study. The metrics produced from the simulated raw survey data and modelled data accurately reflected the underlying mean abundance of the simulated data, but more accurately for the more-common species. Changes in the centre of gravity were detected with high accuracy. The percent of occupied area is generally accurate in its trend when there is a strongly aggregated population, whereas classifying occupied area is more complicated when the population is more dispersed. This preliminary analysis provides the context for analysing distributional metrics at the scale of large marine ecosystems. The spatial distributions of three age groups of plaice were analysed as a case study. The observed northerly and easterly movements of plaice distribution and into deeper waters are interpreted as a response to warming sea temperatures. Consistency in area usage over time was examined to identify those areas critical to plaice. This analysis revealed that the proportion of the population using the ‘critical areas’ (defined as the area holding 75% of the population in 60% of years) was stable over the study period, though some decreases have been noted in recent years. WGECO compared the list of sensitive species identified by WGBYC with the underlying list of species to be monitored under the protection programmes in the EU, using methods reviewed previously in WGECO as well as the sensitivity ratings carried out by IUCN. A resulting list of sensitive species and vulnerable/endangered/critically endangered species is provided. Halibut (Hippoglossus hippoglossus) was the only species suggested by WGBYC which is considered sensitive to fishing. The remaining species, with the exception of the two Hippocampus species, are diadromous species likely to be most sensitive to deterioration of their freshwater habitat. Eleven sensitive species are absent from the list of species to be monitored under protection programmes; WGECO recommends that these 11 should be added to the Commission list to provide information on sensitive and vulnerable species. Further, WGECO recommends that the requirement to return specimens of endangered and critically endangered species to the sea should not be restricted to parts of EU waters and that the status of sensitive species should be monitored on a regular basis, with the aim of providing advice on the adequacy of existing landing restrictions.

Octacalcium phosphate (OCP; Ca8(HPO4)2(PO4)4. 5H2O) is a plausible precursor phase of biological hydroxyapatite, which composites with a number of biologically relevant organic metabolites. Widely used material science physicochemical structure determination techniques successfully characterize the mineral component of these composites but leave details of the structure, and interactions with mineral, of the organic component almost completely obscure. The metabolic linear di-acids succinate (SUC) and adipate (ADI) differentially expand the hydrated (100) layer of OCP. 13C13C correlation (proton driven spin diffusion, PDSD) experiments on OCP composited with (U-13C4)-SUC, and (U13C6)-ADI, show that the two di-acids per unit cell adopt non-centrosymmetric but mutually identical structures. 13C{31P}, rotational echo double resonance (REDOR) shows that one end of each linear di-acid is displaced further from the surface of the apatitic OCP layer relative to the other end. Overall the results indicate two di-acids per unit cell disposed perpendicularly across the OCP hydrated layer with one carboxylate of each di-acid substituting a hydrated surface OCP phosphate group. This study re-affirms the unique advantages of ssNMR in elucidating structural details of organic-inorganic biocomposites, and thereby mechanisms underlying the roles of small metabolites in influencing biomineralization mechanisms and outcomes.

Fibrillar collagens have mechanical and biological roles, providing tissues with both tensile strength and cell binding sites which allow molecular interactions with cell-surface receptors such as integrins. A key question is: how do collagens allow tissue flexibility whilst maintaining well-defined ligand binding sites? Here we show that proline residues in collagen glycine-proline-hydroxyproline (Gly-Pro-Hyp) triplets provide local conformational flexibility, which in turn confers well-defined, low energy molecular compression-extension and bending, by employing two-dimensional 13C-13C correlation NMR spectroscopy on 13C-labelled intact ex vivo bone and in vitro osteoblast extracellular matrix. We also find that the positions of Gly-Pro-Hyp triplets are highly conserved between animal species, and are spatially clustered in the currently-accepted model of molecular ordering in collagen type I fibrils. We propose that the Gly-Pro-Hyp triplets in fibrillar collagens provide fibril “expansion joints” to maintain molecular ordering within the fibril, thereby preserving the structural integrity of ligand binding sites.

Fluorescein is known to exist in three tautomeric forms defined as quinoid, zwitterionic, and lactoid. In the solid state, the quinoid and zwitterionic forms give rise to red and yellow materials, respectively. The lactoid form has not been crystallized pure, although its cocrystal and solvate forms exhibit colors ranging from yellow to green. An explanation for the observed colors of the crystals is found using a combination of UV/Vis spectroscopy and plane-wave DFT calculations. The role of cocrystal coformers in modifying crystal color is also established. Several new crystal structures are determined using a combination of X-ray and electron diffraction, solid-state NMR spectroscopy, and crystal structure prediction (CSP). The protocol presented herein may be used to predict color properties of materials prior to their synthesis.

ICES WGEF recommends that demersal elasmobranchs be managed using spatial proxies 11 for Maximum Sustainable Yield. Here we combine escapement biomass – the percentage of the stock which must be retained each year to conserve it – with maps of predicted abundance of four ray species (cuckoo, thornback, blonde, and spotted), created using Boosted Regression Tree modelling. We then use a Decision Support Tool to generate location and size options for MPAs to protect these stocks, based on the priorities of the various stakeholders, notably the minimisation of fishing effort displacement. Variations of conservation/fishing priorities are simulated, as well as differential priorities for individual species, with a focus on protecting nursery grounds and spawning areas. The result is a complete software package that produces maps of predicted species abundance from limited survey data, allowing disparate stakeholders and policymakers to discuss management options within a mapping interface. No

We consider the problem of regulating the rate of harvesting a natural resource, taking account of the wider system represented by a set of ecological and economic indicators, given differing stakeholder priorities. This requires objective and transparent decision making to show how indicators impinge on the resulting regulation decision. We offer a new scheme for combining indicators, derived from assessing the suitability of lowering versus not lowering the harvest rate based on indicator values relative to their predefined reference levels. Using the practical example of fisheries management under an “ecosystem approach,” we demonstrate how different stakeholder views can be quantitatively represented by weighting sets applied to these comparisons. Using the scheme in an analysis of historical data from the Celtic Sea fisheries, we find great scope for negotiating agreement among disparate stakeholders.

We consider the problem of regulating the rate of harvesting a natural resource, taking account of the wider system represented by a set of ecological and economic indicators, given differing stakeholder priorities. This requires objective and transparent decision making to show how indicators impinge on the resulting regulation decision. We offer a new scheme for combining indicators, derived from assessing the suitability of lowering versus not lowering the harvest rate based on indicator values relative to their predefined reference levels. Using the practical example of fisheries management under an "ecosystem approach," we demonstrate how different stakeholder views can be quantitatively represented by weighting sets applied to these comparisons. Using the scheme in an analysis of historical data from the Celtic Sea fisheries, we find great scope for negotiating agreement among disparate stakeholders.

Skates and rays represent one of the most vulnerable components of fish communities in temperate demersal fisheries such as the Irish Sea. They also tend to be data poor in comparison to commercially exploited teleost fish. Spatial management has been suggested as an important tool to protect these species, but requires an understanding of the abundance distribution, and the relationship the abundance distribution has with the environment at both adult and juvenile life history stages. Here we modelled bottom trawl survey data using delta log-normal boosted regression trees on to derive rays’ spatial abundance, and environmental links. The modelling approach allowed the development of high resolution predictive maps of abundance of four skate and ray species targeted by fishing activity: thornback, spotted, cuckoo and blonde rays. The distributions of these species were driven by a general preference for sand and coarser substrates as well as higher salinities, temperatures and currents speeds. Spatial comparisons between abundance distributions and locations of skate and ray commercial landings indicated that the main hotspots for the investigated species are outside of the main commercial fishing areas and overlap with potential MPAs proposed for wider ecosystem protection. The method offers a useful tool for selecting potential MPA’s to assist the management and conservation of data-poor species. Yes

Cerithidea dohrni (Kobelt, 1890) (Figures 11, 12 A, Q–CC) Cerithium obtusum — Quoy & Gaimard, 1834: 126 –127, Atlas Mollusques pl. 55, figs 18–21 (shell, headfoot, operculum) (not Lamarck, 1822). Cerithium (Cerithidea) kieneri var. dohrni Kobelt, 1890 a: 49 –50, pl. 10, figs 4, 5 (Philippines; lectotype here designated SMF 228131, Fig. 12 Y, Z; 2 paratypes SMF 228132 / 2; Cecalupo 2005: pl. 31, fig. 4). Cerithidea dohrni — Reid et al., 2013: figs 1 (phylogeny), 2 (map). Cerithidea quadrata — Hidalgo, 1904 –1905: 203 (not Sowerby, 1866). Dautzenberg & Fischer, 1906: 410–411 (in part, includes C. quoyii; not Sowerby, 1866). Lozouet, 2008: 286, pl. 88, fig. 5 a, b. (not Sowerby, 1866). Lozouet & Plaziat, 2008: 57, 112, fig. 30, pl. 20, figs 1–3 (not Sowerby, 1866). Cerithidea reidi — Cecalupo, 2005: 316, pl. 31, fig. 4 (not Houbrick, 1986). Cecalupo, 2006: 60, 197, 233 (in part, includes C. reidi, C. anticipata; not Houbrick, 1986). Taxonomic history. This species was introduced as a variety of C. kieneri (= anticipata) by Kobelt (1890 a) and has generally been identified as C. quadrata (= quoyii) since then. It was recognized as distinct by Reid et al. (2013) as a result of molecular analysis. A photograph of only one of the three syntypes in SMF has been seen; this had been marked as ‘lectotype’ by Zilch, but this designation remained unpublished (R. Janssen, pers. comm.) and is confirmed here in order to fix the identity of the taxon. The identity of C. similis Adams, 1855, likewise described from the Philippines, is unknown (see Excluded and Doubtful Species, above). Diagnosis. Shell: spire straight to slightly convex, whorls rounded, periphery angled; aperture flared, anterior canal and apertural projection well developed; 17–35 axial ribs on penultimate whorl, 4–7 after ventrolateral varix; ventrolateral varix an enlarged rib at 190–240 °; 7 spiral cords on spire, 6 cords above periphery on last whorl, giving fine reticulate sculpture; brown with darker spiral cords. Philippines, Sulawesi, Moluccas. COI GenBank AM 932769 –932770, HE 680229, HE 680232. Material examined. 27 lots. Shell (Fig. 12 Q–CC): H = 21.3–43.3 mm. Shape elongated conical (H/B = 2.34–2.50, SH = 3.05–3.45); decollate, 7–9 whorls remaining; spire whorls rounded, suture distinct; spire profile straight to slightly convex; periphery angled; thin to moderate thickness. Adult lip flared, thickened; apertural margin planar in side view; strong anterior projection adjacent to deep notch of anterior canal. Sculpture on spire of straight to slightly curved (opisthocyrt) narrow axial ribs, interspaces 1–2 times width of ribs, 17–35 ribs on penultimate whorl, 4–9 ribs after ventrolateral varix (i.e. about twice as distant as on spire), remaining strong throughout final whorl except for weaker 1 or 2 ribs before aperture, only faint axial wrinkles on base; spire whorls with 7 (8) spiral cords (posterior 2 small), forming small nodules where they cross axial ribs, spiral interspaces 1.5–2 times width of cords, no interpolated threads, 6 (7) spiral cords above periphery on last whorl; overall effect is of fine reticulate sculpture; base with (7) 9–16 spiral ridges, outermost is peripheral cord of same size as primary cords above. Ventrolateral varix a strongly enlarged rib at 190–240 °, forming anteriorly projecting boss at periphery. Surface with fine spiral microstriae on periostracum, strongest on spiral cords. Colour: brown, spiral cords darker brown; aperture pale brown, spiral lines showing through. Animal: Head and base of tentacles pinkish grey with cream spots; anterior half of snout blackish with scattered yellow spots; tentacles pale grey with black rings; sides of foot pale grey, blackish anteriorly, with small yellowish cream spots; sole of foot grey, pinkish towards margin; mantle pale grey with pinkish stripes at margin (corresponding to shell ribs) (based on ethanol-preserved specimens). Quoy & Gaimard (1834) described and illustrated a black snout with one pinkish and two distal transverse yellow bands, a greenish head with brown spots, reddish tentacles with brown rings, the foot yellow spotted with black and the sole violaceous and edged with yellow. Range (Fig. 11): Philippines, Sulawesi, Moluccas. Records: Philippines: Manila Bay, Luzon (NHMUK 20130252; AM C. 33600; USNM 232955); Busuanga (USNM 233012); Abatan R., Bohol (NHMUK 20130255; MNHN); Iwahig, Palawan (USNM 862793); Jolo (USNM 233115). Indonesia: S Bolo Bay, Cape Kawala, Luwuk Peninsula, Sulawesi (ZMB 191378 a); Buru, Moluccas (Quoy & Gaimard, 1834). Quoy & Gaimard (1834) described receiving specimens from the inhabitants of Ile Bourou (Buru, Indonesia) and illustrated the living animal, so that the provenance of this southernmost record cannot be doubted. There are no confirmed records from Borneo and, if correctly identified, the specimen from ‘Borneo’ in Figure 12 A is believed to be wrongly localized. Habitat and ecology. In Bohol (Philippines) Lozouet & Plaziat (2008) found it in a mixed mangrove forest at the mouth of an estuary (salinity 10–31 ppt) and also on the lower trunks of trees and on Nypa, behind the seaward edge of the forest. They noted that is was replaced by C. balteata where the salinity was below 10 ppt. Remarks. This species was recognized as distinct from C. quoyii as a result of molecular analysis (Reid et al. 2013; Fig. 1; see Remarks on C. quoyii). The three species in the C. quoyii group are closely similar, but there are consistent differences in their shells (Table 1). The present species most resembles C. quoyii and the two may prove to be sister taxa when samples of the third member become available. A single tissue sample (no shells available) from eastern Sulawesi showed an uncorrected pairwise distance for COI of 0.075 from three samples from the Philippines, which is high for an intraspecific comparison in this genus and exceeds the minimum distance between recognized species (0.072 for C. anticipata and C. reidi), so further investigation of genetic structure within C. dohrni is desirable. A shell from Buru, also in eastern Indonesia, was illustrated by Quoy & Gaimard (1834: fig. 18); this is narrower than most C. quoyii, has a more rounded periphery and cancellate sculpture; it therefore resembles C. dohrni from the Philippines and is identified accordingly (it could not be confused with the small, slender forms of C. anticipata found further east in New Guinea). These are the only two records of the species from eastern Indonesia. In contrast, C. balteata from the same region is relatively well represented in museum collections, so C. dohrni appears to be scarce there. It is not collected for food in Bohol (Lozouet & Plaziat 2008)., Published as part of Reid, David G., 2014, The genus Cerithidea Swainson, 1840 (Gastropoda: Potamididae) in the Indo-West Pacific region, pp. 1-65 in Zootaxa 3775 (1) on pages 39-41, DOI: 10.11646/zootaxa.3775.1.1, http://zenodo.org/record/285731, {"references":["Kobelt, W. (1890 b) Eine neue Cerithidea. Nachrichtsblatt der deutschen Malakozoologischen Gesellschaft, 1890, 60 - 61.","Quoy, J. R. C. & Gaimard, J. P. (1834) Voyage de Decouvertes de l'Astrolabe. Zoologie. Vol. 3. J. Tastu, Paris, 954 pp.","Lamarck, J. B. P. A. de (1822) Histoire Naturelle de Animaux sans Vertebres. Vols 6 (2), 7. Paris, 232 pp., 711 pp.","Cecalupo, A. (2005) Elenco della famiglia Cerithiidae Ferussac, 1822 (Prosobranchia). Nomenclatura delle specie conosciute o poco note e relativa revisione sistematica. Fossili e Attuali. Vol. I. Quaderni della Civica Stazione Idrobiologica di Milano, 26, 1 - 366.","Reid, D. G., Claremont, M., Smith, L., Shamoto, M., Glaubrecht, M. & Ozawa, T. (2013) Mosaics in the mangroves: allopatric diversification of tree-climbing mudwhelks (Gastropoda: Potamididae: Cerithidea) in the Indo-West Pacific. Biological Journal of the Linnean Society, 110, 564 - 580.","Hidalgo, J. G. (1904 - 1905) Catalogo de los moluscos testaceos de las Islas Filipinas, Jolo y Marianas. I. - Moluscos marinos. Gaceta de Madrid, Madrid, xvi + 408 pp.","Sowerby, G. B. II (1866) Monograph of the genus Cerithidea. In: Reeve, L. A. (Ed.), Conchologia Iconica. Vol. 15. L. A. Reeve, London, pls 1 - 4.","Dautzenberg, P. & Fischer, H. (1906) Liste des mollusques recoltes par M. H. Mansuy en Indo-Chine et au Yunnan et description d'especes nouvelles. Journal de Conchyliologie, 53, 343 - 471.","Lozouet, P. & Plaziat, J. - C. (2008) Mangrove Environments and Molluscs. Abatan River, Bohol and Panglao Islands, Central Philippines. ConchBooks, Hackenheim, 160 pp.","Houbrick, R. S. (1986) Cerithidea reidi, spec. nov., from Western Australia. Veliger, 28, 280 - 286.","Cecalupo, A. (2006) Elenco della famiglia Cerithiidae Ferussac, 1822 (Prosobranchia). Nomenclatura delle specie conosciute o poco note e relativa revisione sistematica. Attuali. Vol. II. Quaderni della Civica Stazione Idrobiologica di Milano, 27, 1 - 373.","Adams, A. (1855) A monograph of Cerithidea, a genus of Mollusca, with descriptions of several new species, from the collection of High Cuming, Esq,: to which are added, descriptions of two new species of Colina, and one of Donax. Proceedings of the Zoological Society of London, 1854, 83 - 87. http: // dx. doi. org / 10.1111 / j. 1469 - 7998.1854. tb 07236. x"]}

Cerithidea sinensis (Philippi, 1848) (Figures 6, 7A–L) Cerithium (Potamides) sinense Philippi, 1848: 19 –20 (China; 2 syntypes MNHN 25694, Fig. 7 A, B). Cerithium sinense — Philippi, 1849: 18, Cerithium pl. 1, fig. 15. Sowerby, 1855: 886, pl. 186, fig. 274. Cerithidea sinensis —H. Adams & A. Adams, 1854: 293. Adams, 1855: 83. Sowerby, 1866: sp. 17, pl. 3, fig. 17. Mabille, 1887: 158. Yen, 1936: 206 –207, pl. 17, fig. 29. Fan, 1989: 37 –38, pl. 5, fig. 15. Ma, 2004: 38, pl. 016D, fig. 13 (map). Kojima et al., 2006: 525 –535, fig. 2 h. Zhang, 2008: 60, fig. Li & Huang, 2012: 152, fig. Reid et al., 2013: figs 1 (phylogeny), 2 (map). Potamides (Cerithidea) sinensis — Tryon, 1887: 164 –165, pl. 34, figs 97, 98. Cerithium (Cerithidea) sinense — Kobelt, 1890 a: 68 –69, pl. 13, figs 8, 9. Cerithidea (Aphanistylus) sinensis —Casto de Elera, 1896: 330. Dautzenberg & Fischer, 1905: 136 –137. Potamides (Aphanistylus) sinensis — Fischer, 1891: 164. Fischer & Dautzenberg, 1904: 416. Aphanistylus moreleti Wattebled, 1886: 59 –60, pl. 4, fig. 5 (Thuan-an, dans la rivière de Hué, Annam [Thuan An, Hue, Vietnam]; 4 syntypes MNHN 25687; 1 syntype MNHN 25688, Fig. 7 J). Potamides (Aphanistylus) moreleti — Fischer, 1891: 164. Fischer & Dautzenberg, 1904: 416. Cerithidea (Aphanistylus) moreleti — Dautzenberg & Fischer, 1905: 138 –139. Cerithidea (Cerithidea) rhizophorarum — Cecalupo, 2006: 147, 239 (not A. Adams, 1855). Taxonomic history. There are two syntypes of Cerithium sinense in MNHN, respectively 18.6 mm (Fig. 7 B) and 27.8 mm (Fig. 7 A) in height; the latter is larger than either the dimension given in the original description (8 lines [17.4 mm]; Philippi 1848) or the figure subsequently published by Philippi (1849: 18 mm). Nevertheless, the two are conspecific and were collected by Largilliert in China (as in the original description), so both are regarded as syntypes. Cerithidea rollei Kobelt, 1890 b was based on shells said to come from China and figures by Kobelt (1890 a) bear a superficial resemblance to C. sinensis. However, this is a synonym of Cerithideopsis scalariformis from the western Atlantic (see Excluded and doubtful species). Tryon (1887) first suggested the synonymy of C. moreleti with C. sinensis. Fischer (1891) listed both, but Dautzenberg & Fischer (1905) remarked on their close similarity, while provisionally retaining them as separate species. The records of C. charbonnieri from northern Vietnam in early French literature probably apply to C. sinensis (e.g. Mabille, 1887; Crosse & Fischer, 1890; see Range of C. charbonnieri). Diagnosis. Shell: delicate, small, narrow, straight-sided, periphery rounded; sinuous apertural margin, weak anterior canal; 14–19 rounded axial ribs on penultimate whorl, ventrolateral varix often absent; no spiral sculpture above periphery; weakly lined pattern. China, northern Vietnam. COI GenBank HE 680295 – HE 680297. Material examined. 25 lots. Shell (Fig. 7 A–L): H = 16.7–28.6 mm. Shape narrowly elongate (H/B = 2.25–2.78, SH = 3.01–3.45); decollate, 6–7 whorls remaining; spire whorls rounded, suture distinct; spire profile almost straight; periphery rounded; delicate. Adult lip only moderately thickened and flared; rarely, 1–3 previous lips on final whorl (Fig. 7 I); apertural margin sinuous in side view; weak anterior projection adjacent to shallow notch of canal. Sculpture on spire of straight to slightly opisthocline axial ribs, becoming slightly curved (opisthocyrt) and weak on final whorl, ribs rounded, ribs and interspaces of equal width, 14–19 ribs on penultimate whorl; spiral sculpture absent above periphery; base with 8–10 striae, of which 2 at periphery are slightly more prominent. Ventrolateral varix usually absent or indistinct; if present, at 90–140 ° (1 spec. with 2 varices, at 160 and 200 °). Surface with very faint spiral microstriae on periostracum; shiny where periostracum worn away. Colour: fawn to pale brown; 0–2 narrow brown bands above periphery; almost always a strong brown band just below periphery, sometimes another adjacent to columella; periostracum olive brown; bands visible within aperture. Animal: Head and foot cream with grey mottling; snout with two transverse black bands, tip yellow; tentacles yellow at base with black stripe across eye; edge of foot mottled yellow; mantle edge pale grey (based on ethanolpreserved specimens). Range (Fig. 6): Asian mainland from Yellow Sea to N Vietnam. Records: China: Liaoning to Jiangsu (Ma, 2004); Shanghai (NHMUK 1867.5.18.14); Ningbo (ANSP 18093); Un-Long, Kowloon, Hong Kong (USNM 516424). Vietnam: Haiphong (NHMUK 1893.12.8.120; USNM 207877; MNHN); Nam Dinh, Tonkin (MNHN); Da Nang (NHMUK 1901.12.12.175; MNHN; ZMB). Ma (2004) reported this species to be restricted to northern China, but records are here extended south to Vietnam. Pilsbry (1895) recorded it from the Yaeyama Islands at the southern end of the Ryukyu chain, but all other records are from continental coasts, so this is considered unlikely and could refer to small specimens of C. moerchii. Habitat and ecology. Yen (1936) reported it to be very common in brackish water at a river mouth (Shantung Peninsula). The type locality of Aphanistylus moreleti is the Hue River, where Wattebled (1886: 59) noted that it was “fairly common”. Records from southern China and Vietnam are scarce, suggesting that it may be less frequent there. It has not been recorded if this species climbs on vegetation, but its range in the north of China lies beyond that of mangrove trees. Remarks. This species is well known in the Chinese literature, but not elsewhere, and is scarce in western museum collections. Specimens from the extreme south of the range at Da Nang and Hue in northern Vietnam are smaller (Fig. 7 J, K) than the typical form from China (Fig. 7 A–D), but the two appear to integrade at Haiphong in northern Vietnam (Fig. 7 E–L). The southern form was recognized as C. moreleti by some early twentieth century authors (see Synonymy). Apart from size, there are no consistent differences in the shells of northern and southern forms, but their conspecificity needs to be tested with molecular data. There is geographical overlap with C. tonkiniana over much of the range. Dautzenberg & Fischer (1905) considered tonkiniana to be a variety of C. sinensis and the two names have been confused on some museum samples. The shell of C. tonkiniana is larger (to 38.1 mm), more solid, has a strong ventrolateral varix and 5 weak spiral ridges above the periphery (if not eroded; Fig. 13 N–Y); the ventrolateral varix and spiral ridges are absent in C. sinensis. It is unclear if the two differ ecologically, but C. sinensis possibly occurs in habitats of lower salinity. Comparison with the two most closely related congeners, C. balteata and C. charbonnieri, is discussed in the Remarks on C. balteata. Another potamidid that is broadly sympatric with C. sinensis is Cerithideopsis largillierti. The latter species is not decollate, has more rounded whorls, the axial ribs are more numerous (20–26 on penultimate whorl), the basal cords are more pronounced, the aperture is more strongly sinuous in side view and is not flared, the periostracum is thicker with strong, slightly bristly striae and the living animal is black with yellow tip to the snout and yellow tentacle bases., Published as part of Reid, David G., 2014, The genus Cerithidea Swainson, 1840 (Gastropoda: Potamididae) in the Indo-West Pacific region, pp. 1-65 in Zootaxa 3775 (1) on pages 23-25, DOI: 10.11646/zootaxa.3775.1.1, http://zenodo.org/record/285731, {"references":["Philippi, R. A. (1848) testaceorum novorum centuria (continuatio). Zeitschrift fur Malakozoologie, 1848, 17 - 27.","Philippi, R. A. (1849) Abbildungen und Beschreibungen neuer oder wenig gekannter Conchylien. Vol. 3. Part 4. Theodor Fischer, Cassel, 28 pp.","Sowerby, G. B. II (1855) Monograph of the genus Cerithium, Adanson. In: Sowerby, G. B. II. (Ed.), Thesaurus Conchyliorum, or Figures and Descriptions of Recent Shells. Vol. 2. part 16. Sowerby, London, pp. 847 - 899.","Adams, A. (1855) A monograph of Cerithidea, a genus of Mollusca, with descriptions of several new species, from the collection of High Cuming, Esq,: to which are added, descriptions of two new species of Colina, and one of Donax. Proceedings of the Zoological Society of London, 1854, 83 - 87. http: // dx. doi. org / 10.1111 / j. 1469 - 7998.1854. tb 07236. x","Sowerby, G. B. II (1866) Monograph of the genus Cerithidea. In: Reeve, L. A. (Ed.), Conchologia Iconica. Vol. 15. L. A. Reeve, London, pls 1 - 4.","Mabille, J. (1887) Sur quelques mollusques du Tonkin. Bulletins de la Societe Malacologique de France, 4, 73 - 164.","Yen, T. C. (1936) The marine gastropods of Shantung Peninsula. Contributions from the Institute of Zoology, National Academy of Peiping, 3, 165 - 255.","Fan, C. (Ed.) (1989) Mollusca of Huanghai and Bohai. Nongye Chubanshe, Agricultural Publishing House, Beijing, 309 pp.","Ma, X. (2004) Order Mesogastropoda. In: Qi, Z. (Ed.), Seashells of China. China Ocean Press, Beijing, pp. 31 - 81.","Kojima, S., Kamimura, S., Iijima, A., Kimura, T., Kurozumi, T. & Furota, T. (2006) Molecular phylogeny and population structure of tideland snails in the genus Cerithidea around Japan. Marine Biology, 149, 525 - 535. http: // dx. doi. org / 10.1007 / s 00227 - 005 - 0183 - 2","Zhang, S. (2008) Atlas of Marine Mollusks in China. Hai yang chu ban she, Beijing, iv + 383 pp.","Li, R. & Huang, Z. (2012) Class Gastropoda. In: Huang, Z. & Liu, M. (Eds.), The Living Species and their Illustrations in China's Seas (Part II). An Illustrated Guide to Species in China's Sea. Vol. 4. Ocean Press, Beijing, pp. 102 - 279.","Reid, D. G., Claremont, M., Smith, L., Shamoto, M., Glaubrecht, M. & Ozawa, T. (2013) Mosaics in the mangroves: allopatric diversification of tree-climbing mudwhelks (Gastropoda: Potamididae: Cerithidea) in the Indo-West Pacific. Biological Journal of the Linnean Society, 110, 564 - 580.","Tryon, G. W. (1887) Manual of Conchology. Vol. 9. Tryon, Philadelphia, 488 pp.","Kobelt, W. (1890 b) Eine neue Cerithidea. Nachrichtsblatt der deutschen Malakozoologischen Gesellschaft, 1890, 60 - 61.","Casto de Elera, R. P. (1896) Catalogo Sistematico de Toda la Fauna de Filipinas, Vol. 3. Moluscos y Radiados. Colegio de Santo Tomas, Manila, 942 + lxiv pp.","Dautzenberg, P. & Fischer, H. (1905) Liste des mollusques recoltes par M. le Capitaine de Fregate Blaise au Tonkin, et description d'especes nouvelles. Journal de Conchyliologie, 53, 85 - 234.","Fischer, P. (1891) Catalogue et distribution geographique des mollusques terrestres, fluviatiles et marins d'une partie de l'Indo- Chine (Siam, Laos, Cambodge, Annam, Tonkin). Bulletin de la Societe d'Histoire Naturelle d'Autun, 4, 84 - 276.","Fischer, H. & Dautzenberg, P. (1904) Catalogue des mollusqes terrestres et fluviatiles de l'Indo-Chine orientale cites jusqu'a ce jour. In: Pavie, A. (Ed.), Mission Pavie Indo-Chine 1879 - 1895. Etudes Diverses. III. Recherches sur l'Histoire Naturelle de l'Indo-Chine Orientale. E. Leroux, Paris, pp. 390 - 450.","Wattebled, G. (1886) Description de mollusques inedits de l'Annam. Recolte du capitaine Dorr aux environs de Hue. Journal de Conchyliologie, 34, 54 - 71.","Cecalupo, A. (2006) Elenco della famiglia Cerithiidae Ferussac, 1822 (Prosobranchia). Nomenclatura delle specie conosciute o poco note e relativa revisione sistematica. Attuali. Vol. II. Quaderni della Civica Stazione Idrobiologica di Milano, 27, 1 - 373.","Crosse, H. & Fischer, P. (1890) Mollusques marins de la Baie d'Halong (Tonkin). Journal de Conchyliologie, 38, 14 - 19.","Pilsbry, H. A. (1895) Catalogue of the Marine Molluscs of Japan, with Descriptions of New Species and Notes on Others Collected by F. Stearns. F. Stearns, Detroit, viii + 196 pp."]}

Cerithidea tonkiniana Mabille, 1887 (Figures 2 F, 13 N–Y, 14 A) Cerithidea tonkiniana Mabille, 1887: 158 –159 (Tonkin [N Vietnam]; 3 syntypes MNHN 25690, Fig. 13 Q]. Reid et al., 2013: figs 1 (phylogeny), 2 (map). Potamides (Cerithidea ? ) tonkinianus — Fischer, 1891: 163. Potamides (Cerithidea) tonkinianus — Fischer & Dautzenberg, 1904: 416. Cerithidea (Cerithidea) sinensis var. tonkiniana — Dautzenberg & Fischer, 1905: 137 –138. Cerithidea ornata — Kuroda & Habe, 1952: 44. Habe & Kosuge, 1967: 34, pl. 13, fig. 17. Habe, 1968: 38, pl. 11, fig. 24. Higo, 1973: 59. Wells, 1985: 145, pl. 1 G. Choe, 1992: 310 –311, pl. 65, fig. 103 a, b. Higo & Goto, 1993: 105. Higo, Callomon & Goto, 1999: 87. Lee & Min, 2002: 103. Ma, 2004: 38, pl. 016A (not 016C). Kojima et al., 2006: 525 –535. Reid et al., 2008: 680 –699 (in part, includes C. balteata). Zhang, 2008: 61, fig. (All not Sowerby, 1855). Cerithidea (Cerithidea) ornata — Oyama, 1959: figs 7, 14. Fukuda, 1996: 21, pl. 3, fig. 4. Fukuda, 2000: 106 –110, 135, fig. 5.4 b (shell), fig. 5.5 b (living animal). Hasegawa, 2000: 133, pl. 66, fig. 2. Hong, Choi & Tsutsumi, 2010: 258, fig. 4 b. Fukuda & Kimura, 2012: 29, textfigs a (living animal), b, c (shell). (All not Sowerby, 1855). Cerithidea morchii — Lee & Min, 2002: 103 (not Sowerby, 1855). Min et al., 2004: 129, fig. 220 (not Sowerby, 1855). Cerithidea (Cerithidea) weyersi — Thach, 2007: 60, pl. 9, figs 173, 181 (not Dautzenberg, 1899). Cerithidea sinensis — Li & Xu, 2013: 452 –453, figs 8 (shell), 9 (radula) (not Philippi, 1848). Taxonomic history. The original description by Mabille (1887) was rather vague and the species was not illustrated. For these reasons the name has not been used since the early twentieth century, until recently resurrected by Reid et al. (2013). Dautzenberg & Fischer (1905) considered it to be a variety of C. sinensis. The species has frequently been described and figured in the recent literature from Japan, China, Korea and Vietnam, although always misidentified. It has usually been named C. ornata (a synonym of the superficially similar C. balteata), following the identification by early Japanese workers (Kuroda & Habe 1952; Oyama 1959). Diagnosis. Shell: periphery rounded or slightly angled; aperture thickened and flared; 11–25 rounded axial ribs on penultimate whorl, 4–11 ribs after ventrolateral varix; ventrolateral varix a strong rib at 180–220 (250)°; 5 weak spiral ridges above periphery (especially on spire if not eroded); banded or lined pattern. S Japan, S Korea, S China, N Vietnam. COI GenBank AM 932766, AM 932767, HE 680220 –680224. Material examined. 15 lots. Shell (Fig. 13 N–Y): H = 23.0– 38.1 mm. Shape elongated conical (H/B = 2.04–2.31, SH = 2.59–2.99); decollate, 6–7 whorls remaining; spire whorls moderately rounded, suture distinct; spire profile mostly straight, but concave in apical whorls that are lost in adults and becoming convex at final whorl; periphery rounded to slightly angled; thin to moderate thickness. Adult lip flared, slightly thickened; rarely a previous lip on final whorl; apertural margin slightly sinuous in side view; moderate anterior projection adjacent to canal. Sculpture on spire of straight to slightly opisthocline axial ribs, becoming slightly curved (opisthocyrt) on final 1–2 whorls, ribs prominent, rounded, interspaces 1–1.5 times width of ribs, 11–25 ribs on penultimate whorl; axial ribs becoming more distant but usually remaining strong after ventrolateral varix, numbering 4–11; spiral sculpture only visible on spire whorls if well preserved, 5 weak spiral ridges on axial ribs only, plus strong outermost basal cord usually visible in suture; traces of 5 spiral ridges sometimes present above periphery on final whorl, but only in interspaces between axial ribs; base with 9–13 striae, of which outermost is largest and may form a weak peripheral keel, below which axial ribs do not continue. Ventrolateral varix a thickened rib at 180–220 (250)°. Surface with fine spiral microstriae on periostracum; ribs dull when worn. Colour: pale brown with dark spiral lines and bands; basic pattern is brown band on base, a line just below periphery and 4 bands at and above periphery, but these may be lost or fused; even in darkest shells there is a pale band at shoulder and below periphery; bands show through in aperture. Animal (Fig. 2 F): Head grey with cream spots; anterior half of snout blackish with yellow spots arranged in 2–3 transverse bands; tentacles grey with black rings and black base; sides of foot blackish; sole of foot grey; mantle pale grey (ethanol-preserved specimens similar). Range (Fig. 14 A): S Japan, S Korea, S China, Vietnam. Records: Japan: Tosa Prov., Shikoku (ANSP 191129); Ukidono-kobashi, Yorimo R., Touruta-shinden, Usa, Oita Pref., Kyushu (NHMUK 20070383); Hikawa R., Yatsushiro-gun, Kumamoto Pref., Kyushu (NHMUK 20070378). S Korea: Suncheon Bay (Hong, Choi & Tsutsumi 2010). China: Zhejiang to Guangxi (Qi 2004); Tolo Harbour, Hong Kong (NHMUK 20130261; USNM 858380). Vietnam: Quikim Brackish Water Research Station, Hai Phong (NHMUK 20130260); Ha Long Bay (RMNH); Long Hai (Thach 2007, as C. weyersi). Habitat and ecology. On mangrove trunks and branches and on mud and stones under mangroves (Hong Kong); intertidal mudflat with Phragmites reeds at river mouth (Japan); brackish salt marsh at edge of shrimp ponds (northern Vietnam). In Hong Kong, Wells (1985) reported it from mangroves at upper tidal levels, attached by mucus to trees, and described it as rare, except at Ting Kok. McMahon (1985) likewise observed that it was restricted to the upper parts of the mangrove belt, above the mean tide mark; it was most common at the landward edge and found above 1.0 m on trunks and branches, attached by mucus when dry. Hong et al. (2010) found it on mud flats among Suaeda plants in Korea. Remarks. The shell of this species closely resembles that of C. balteata from the Philippines, Indonesia and New Guinea, and it has usually been identified as ‘ C. ornata ’ (= C. balteata). Molecular data show that the two are distinct and belong to separate clades (Reid et al. 2008, 2013). The most useful characters for separation are the broader shape of C. tonkiniana with stronger and more numerous (4–11, cf. 0–6 in C. balteata) axial ribs continuing after the ventrolateral varix; the frequent position of the ventrolateral varix at about 180 ° (usually about 270 ° in C. balteata); and the weak spiral ribs on the spire whorls (absent in C. balteata). However, the great variability of C. balteata across its wide range and the often eroded condition of the shells make morphological diagnosis difficult. Cerithidea tonkiniana is sympatric over much of its range in China and northern Vietnam with C. sinensis, with a somewhat similar shell (see Remarks on C. sinensis for discrimination). Its range also overlaps broadly with that of another potamidid, Cerithideopsis largillierti. Although superficially similar, the latter species is not decollate, lacks a distinct ventrolateral varix, the aperture is sinuous in side view and is not flared, the periostracum is thicker with strong slightly bristly striae, the whorls are more rounded and the axial ribs are often more numerous (20–26 on penultimate whorl); the living animal is black with yellow snout tip and tentacle bases, and does not climb onto vegetation. In molecular analyses C. tonkiniana forms a clade with C. moerchii (Reid et al., 2008, 2013; Fig. 1) although the sister species of the latter is probably the unsampled C. rhizophorarum (see Remarks on that species). Cerithidea tonkiniana and C. moerchii are broadly sympatric from southern Japan to Vietnam, but are readily distinguished by the much stronger spiral ridges of C. moerchii (Fig. 15 J–AA). The larval development has not been explicitly reported, but Kojima et al. (2006) claimed a ‘planktotrophic stage’ of 12–20 days (quoting Kimura et al. 2002, who mentioned planktonic larvae in ‘ C. ornata ’, and unpublished work). This species is listed as ‘endangered’ in Japan (Fukuda 1996; Fukuda & Kimura 2012) as a consequence of threats to its habitat and its narrow range in the country., Published as part of Reid, David G., 2014, The genus Cerithidea Swainson, 1840 (Gastropoda: Potamididae) in the Indo-West Pacific region, pp. 1-65 in Zootaxa 3775 (1) on pages 47-48, DOI: 10.11646/zootaxa.3775.1.1, http://zenodo.org/record/285731, {"references":["Mabille, J. (1887) Sur quelques mollusques du Tonkin. Bulletins de la Societe Malacologique de France, 4, 73 - 164.","Reid, D. G., Claremont, M., Smith, L., Shamoto, M., Glaubrecht, M. & Ozawa, T. (2013) Mosaics in the mangroves: allopatric diversification of tree-climbing mudwhelks (Gastropoda: Potamididae: Cerithidea) in the Indo-West Pacific. Biological Journal of the Linnean Society, 110, 564 - 580.","Fischer, P. (1891) Catalogue et distribution geographique des mollusques terrestres, fluviatiles et marins d'une partie de l'Indo- Chine (Siam, Laos, Cambodge, Annam, Tonkin). Bulletin de la Societe d'Histoire Naturelle d'Autun, 4, 84 - 276.","Fischer, H. & Dautzenberg, P. (1904) Catalogue des mollusqes terrestres et fluviatiles de l'Indo-Chine orientale cites jusqu'a ce jour. In: Pavie, A. (Ed.), Mission Pavie Indo-Chine 1879 - 1895. Etudes Diverses. III. Recherches sur l'Histoire Naturelle de l'Indo-Chine Orientale. E. Leroux, Paris, pp. 390 - 450.","Dautzenberg, P. & Fischer, H. (1905) Liste des mollusques recoltes par M. le Capitaine de Fregate Blaise au Tonkin, et description d'especes nouvelles. Journal de Conchyliologie, 53, 85 - 234.","Kuroda, T. & Habe, T. (1952) Check List and Bibliography of the Recent Marine Mollusca of Japan. L. W. Stach, Tokyo, 210 pp.","Habe, T. & Kosuge, S. (1967) The Standard Book of Japanese Shells in Colour, Vol. III. Hoikusha, Osaka, xviii + 223 pp.","Habe, T. (1968) Shells of the Western Pacific in Color. Vol. 2. 2 nd ed. Hoikusha, Osaka, 233 pp.","Higo, S. (1973) A Catalogue of Molluscan Fauna of the Japanese Islands and the Adjacent Area. Isahaya, 397 + 61 pp.","Wells, F. E. (1985) The Potamididae (Mollusca: Gastropoda) of Hong Kong, with an examination of habitat segregation in a small mangrove system. In: Morton, B. & Dudgeon, D. (Eds) Proceedings of the Second International Workshop on the Malacofauna of Hong Kong and Southern China, Hong Kong. Hong Kong University Press, Hong Kong, pp. 139 - 154.","Choe, B. L. (1992) Mollusca (II). Illustrated Encyclopaedia of Fauna and Flora of Korea. Vol. 33. Ministry of Education, Republic of Korea, 860 pp.","Higo, S. & Goto, Y. (1993) A Systematic List of Molluscan Shells from the Japanese Islands and the Adjacent Areas. Eru Malacological Publisher, Osaka, 3 + 22 + 693 + 13 + 149 pp.","Higo, S., Callomon, P. & Goto, Y. (1999) Catalogue and Bibliography of the Marine Shell-Bearing Mollusca of Japan. Elle Scientific Publications, Osaka, 749 pp.","Lee, J. - S. & Min, D. - K. (2002) A catalogue of molluscan fauna in Korea. Korean Journal of Malacology, 18, 93 - 217.","Ma, X. (2004) Order Mesogastropoda. In: Qi, Z. (Ed.), Seashells of China. China Ocean Press, Beijing, pp. 31 - 81.","Kojima, S., Kamimura, S., Iijima, A., Kimura, T., Kurozumi, T. & Furota, T. (2006) Molecular phylogeny and population structure of tideland snails in the genus Cerithidea around Japan. Marine Biology, 149, 525 - 535. http: // dx. doi. org / 10.1007 / s 00227 - 005 - 0183 - 2","Reid, D. G., Dyal, P., Lozouet, P., Glaubrecht, M. & Williams, S. T. (2008) Mudwhelks and mangroves: the evolutionary history of an ecological association (Gastropoda: Potamididae). Molecular Phylogenetics and Evolution, 47, 680 - 699. http: // dx. doi. org / 10.1016 / j. ympev. 2008.01.003","Zhang, S. (2008) Atlas of Marine Mollusks in China. Hai yang chu ban she, Beijing, iv + 383 pp.","Sowerby, G. B. II (1855) Monograph of the genus Cerithium, Adanson. In: Sowerby, G. B. II. (Ed.), Thesaurus Conchyliorum, or Figures and Descriptions of Recent Shells. Vol. 2. part 16. Sowerby, London, pp. 847 - 899.","Oyama, K. (1959) Cerithidea. In: Oyama, K. (Ed.), The Molluscan Shells, Vol. 2. Resources Exploitation Institute, Tokyo, unpaginated.","Fukuda, H. (1996) Gastropoda. In: Hanawa, S. & Sakuma, H. (Eds.), Present Status of Estuarine Locales and Benthic Invertebrates Occurring in Estuarine Environment in Japan. WWF Japan Science Report, 3, 11 - 63.","Fukuda, H. (2000) Gastropods I: general view. In: Sato, M. (Ed.), Life in Ariake Sea: Biodiversity in Tidal Flats and Estuaries. Kaiyu-sha, Tokyo, pp. 213 - 252.","Hasegawa, K. (2000) Family Potamididae. In: Okutani, T., (Ed.), Marine Mollusks in Japan. Tokai University Press, Tokyo, pp. 133 - 134.","Hong, J. - S., Choi, J. - W. & Tsutsumi, H. (2010) Concluding remarks on the joint survey of macrobenthic fauna on Suncheon Tidal Flats by the participants of \" Korea and Japan Joint Symposium on Biology of Tidal Flats 2009 \". Plankton & Benthos Research, 5 (suppl.), 255 - 263. http: // dx. doi. org / 10.3800 / pbr. 5.255","Min, D. - K., Lee, J. - S., Koh, D. - B. & Je, J. - G. (2004) Mollusks in Korea. Min Molluscan Research Institute, Seoul, 566 pp.","Thach, N. N. (2007) Recently Collected Shells of Vietnam. L'Informatore Piceno, Ancona, Italy, 384 pp.","Dautzenberg, P. (1899) Contribution a la faune malacologique de Sumatra (recoltes de M. J. - L. Weyers). Annales de la Societe Royale Malacologique de Belgique, 34, 1 - 26.","Li, Y. & Xu, R. (2013) Brackish water snails from Qi'ao-Dan'gan Island in the Pearl River estuary, China. Turkish Journal of Zoology, 37, 449 - 457.","Philippi, R. A. (1848) testaceorum novorum centuria (continuatio). Zeitschrift fur Malakozoologie, 1848, 17 - 27.","McMahon, R. F. (1985) Interspecific relationships between morphometric parameters and the vertical distribution patterns of seven species of turbinate gastropods on mangrove trees in Hong Kong. In: Morton, B. & Dudgeon, D. (Eds.), Proceedings of the Second International Workshop on the Malacofauna of Hong Kong and Southern China, Hong Kong. Hong Kong University Press, Hong Kong, pp. 199 - 215.","Kimura, T., Fujioka, E., Kimura, S. & Aoki, S. (2002) A comparative morphology of eggs and larvae of nine gastropods in tidal flat and marsh of the reed. Venus, 61, 114 - 115. [in Japanese]"]}

Cerithidea decollata (Linnaeus, 1767) (Figures 9, 13A–M) Murex decollatus Linnaeus, 1767: 1226 (no locality; neotype, here designated, NHMUK 20130229, SE Pemba Bay, Cabo Delgado Prov., Mozambique; Fig. 13 J–M). Gmelin, 1791: 3563. Dillwyn, 1817: 759. Hanley, 1855: 311. Dodge, 1957: 204 –206. Cerithium decollatum —Bruguière, 1792: 501. Lamarck, 1822: 71. Kiener, 1841 –1842: 96–97, pl. 28, fig. 2. Deshayes, 1843: 294. Sowerby, 1855: 886, pl. 186, fig. 276. Cerithidea decollata — Gray, 1847: 154. H. Adams & A. Adams, 1854: 292–293, pl. 31, fig. 2 a (operculum). Adams, 1855: 83. Troschel, 1858: 147, pl. 12, fig. 4 (radula). Sowerby, 1866: sp. 14, pl. 2, fig. 14 a, b. Haas, 1929: 427. Barnard, 1963: 140, fig. 25 d (radula). Macnae, 1968: figs 49, 55. Fischer-Piette & Vukadinovic, 1973: 361 –362, fig. 15, 16 (map). Kilburn & Rippey, 1982: 53, pl. 10, fig. 17. Brown, 1994: 143, fig. 68 b. Reid et al., 2008: 680 –699, figs 1, 2 (phylogeny). Reid et al., 2013: figs 1 (phylogeny), 2 (map). Cerithium (Potamides) decollatum — Krauss, 1848: 108. Potamides (Cerithidea) decollatus —von Martens, 1880: 282. Tryon, 1887: 161, pl. 32, fig. 54 (as decollata). von Martens, 1897 a: 188. von Martens, 1897 b: 266. Dautzenberg, 1923: 48. Dautzenberg, 1929: 489. Cerithium (Cerithidea) decollatum — Kobelt, 1890 a: 44 –45, pl. 9, figs 6, 7. Cerithidea (Cerithidea) decollata —Thiele, 1929: 206–207, fig. 203 (radula). Starmühlner, 1969: 243 –244. Cecalupo, 2006: 57, 194– 195, pl. 57, fig. 8 a–c. ? Turbo pulcher Dillwyn, 1817: 855 (refers to Lister 1688: pl. 588, fig. 50; coasts of the West India Islands; in part, includes Epitoniidae). Cerithidea (Pirenella) decollata (Lamarck) — Boettger, 1890 b: 97 –98 (not Melania decollata Lamarck, 1822). Cerithidea (Cerithidea) rhizophorarum — Cecalupo, 2006: 136, 234, pl. 57, figs 7, 9– 11 (in part, includes C. rhizophorarum, C. moerchii, C. weyersi; not A. Adams, 1855) Taxonomic history. Although this is now such a well-known species, there has been some uncertainty about its identification as Murex decollatus Linnaeus, 1767. The evidence has been discussed in detail by Dodge (1957). Briefly, Linnaeus’ (1767) description was typically short, quoted no figure, gave no locality and mentioned only the Museo de Geer as the source of the specimen(s). No specimens are present in the Linnean Collection in London (Hanley 1855; Dodge 1957). According to Dance (1986), the De Geer collection was in Stockholm, but no material now survives in the Swedish Museum of Natural History (A. Warén, pers. comm.). Hanley (1855: 311) considered Linnaeus’ description to be “wholly insufficient for the purpose of definition”, but nevertheless concluded that “it is not desirable to suggest another hypothetical representative” to replace the established concept that he attributed to Bruguière (1789 –1792). Following the original description by Linnaeus (1767), Gmelin (1791) added no further details and the first detailed written description of the taxon was indeed that of Bruguière (1789 –1792). This latter account was evidently based on a series of specimens and mentioned the fine spiral striae between the longitudinal ribs, the ventrolateral varix, and colour of pale ribs and two brown spiral lines; although it gave no locality it is, however, sufficient to confirm Bruguière’s identification. This concept was followed by other early French authors (Lamarck 1822; Kiener 1841 –1842; Deshayes 1843), of whom both Deshayes and Kiener attributed the specific name to Bruguière, presumably considering Linnaeus’ description inadequate for recognition. The first figures were provided by Kiener (1841 –1842) and Sowerby (1855). Subsequently, some other authors have continued to attribute the name C. decollata to Bruguière (von Martens 1880, 1897a, b; Haas 1929; Starmühlner 1969; Fischer- Piette & Vukadinovic 1973). Quoting Hanley (1855), von Martens (1897 a: 189) denied that Murex decollatus Linnaeus, 1767 was the same species as that described by Bruguière, instead proposing that the Linnean taxon was probably the South American “ Melania atra Richard (truncata Lam.)”. Nevertheless, the majority of authors have accepted the identification of the present species as Murex decollatus Linnaeus, 1767. Reviewing the evidence, Dodge (1957: 205–206) considered Linnaeus’ (1767) description “ample and fairly clear” and dismissed Hanley’s objection as “trivial”. It does, however, seem desirable that the identity of the species should be established unequivocally with a type specimen and therefore a neotype is designated here. Turbo pulcher Dillwyn, 1817 is a doubtful synonym. The author referred to Gmelin (1791: 3603–3604), Schröter (1784: 108) and Lister (1688: pl. 588, fig. 50). Deshayes (1843) was the first to list this species in the synonymy of C. decollata, but with a query, and this was followed by Adams (1855), Sowerby (1855, 1866) and Tryon (1887). Lister’s (1688) figure could indeed represent C. decollata, the shape of the aperture and longitudinal ribs being characteristic; the engraving also suggests a possible ventrolateral varix, although whether the apex is decollate is arguable. Both Gmelin (1791) and Schröter (1784) identified the figure as a shell allied to Turbo clathrus Linnaeus, 1758, a member of the Epitoniidae, and their written descriptions apply to epitoniids also. Dillwyn’s (1817) description is not precise and it appears that he was simply quoting the earlier authors and not describing a specimen. It is not clear why he gave the locality as the West Indies. There has been some confusion with Melania decollata Lamarck, 1822. This can be traced to Boettger (1890 b), who included references to that species and to C. decollata in the synonymy of his “ Cerithidea (Pirenella) decollata (Lmk.) ”, while describing C. decollata. References to Melania decollata were subsequently included in synonymies of C. decollata compiled by Haas (1929), Starmühlner (1969) and Fischer-Piette & Vukadinovic (1973). Kobelt (1893) introduced the taxa Cerithium (Cerithidea) inaequisculptum and Cerithium (Cerithidea) natalense, and Sowerby (1855) described Cerithium rissoideum. All three were said to originate from Natal, but all are species of Cerithideopsis from the western Atlantic (see Excluded and doubtful species, above). Diagnosis. Shell: spire convex, whorls moderately rounded, periphery rounded; aperture flared, apertural projection well developed; 18–41 axial ribs on penultimate whorl, 4–12 weakening ribs after ventrolateral varix; ventrolateral varix an enlarged rib at 170–240 °; 13–18 spiral ridges develop on final 1–4 whorls, 12–17 ridges above periphery on last whorl; brown with darker spiral lines and bands. S and E Africa, Madagascar. COI GenBank AM 932763, AM 932764, HE 680207. Material examined. 54 lots. Shell (Fig. 13 A–M): H = 19.5–37.7 mm. Shape elongated conical (H/B = 1.84–2.28, SH = 2.41–2.84); decollate, 5–6 whorls remaining; spire whorls moderately rounded, suture distinct; spire profile convex; periphery rounded; thickness moderate. Adult lip flared, thickened; apertural margin planar in side view; weak anterior projection adjacent to notch of anterior canal; columella slightly detached. Sculpture on spire of straight axial ribs, becoming slightly curved (opisthocyrt) on last 2 whorls, ribs and interspaces of equal width, 18–41 ribs on penultimate whorl, 4–12 ribs after ventrolateral varix, weakening on final 0.25 whorl (final 0.5 whorl often regrown after damage, axial ribs then lacking); only faint axial wrinkles on base; spiral ridges develop only on final 1–4 whorls, sometimes faint and visible only anteriorly and in interspaces between axial ribs, 13–18 fine spiral ridges visible on penultimate and some earlier whorls, separated by incised lines, peripheral cord often visible in suture; 12–17 fine spiral ridges above periphery on last whorl; base with 11–16 fine spiral ridges of same size as those above periphery (on earlier whorls peripheral cord is slightly enlarged). Ventrolateral varix a moderately enlarged rib at 170–200 (240)°. Surface with fine spiral microstriae on periostracum. Colour: pale brown, with 5 dark brown spiral bands (2 above, 1 at periphery, 2 on base), posterior 2 often fused to form broad band with paler zone at suture and periphery; aperture brown, spiral bands showing through. Animal: Head pinkish grey with cream spots; anterior half of snout blackish with transverse black line at tip; tentacles grey with black rings and black base; sides of foot blackish; sole of foot grey; mantle pale pinkish grey (based on ethanol-preserved specimens). Range (Fig. 9): S and E Africa, Madagascar. Kenya: Mombasa (NHMUK; USNM 215238). Tanzania: Chukwani, Zanzibar (ANSP 214825); Lindi (USNM 595347; ZMB). Mozambique: SE Pemba Bay, Cabo Delgado Prov. (NHMUK 20060485). South Africa: Richards Bay, Kwazulu Natal (AM); Zwartkops R., Port Elizabeth, Cape Prov. (NHMUK 20130257); Knysna (Macnae 1963; Hodgson & Dickens 2012). Madagascar: Amboaniva (Fischer-Piette & Vukadinovic 1973); Ambataloaka, SW Nossi Bé (USNM 633348; ANSP 258481); Ambongo (MNHN); Toliara (NHMUK 20130258). There are several old and incompletely localized records from Réunion and Mauritius (MNHN, USNM), but these are considered unreliable, because the species was not recorded from Mauritius by Viader (1937) or from Réunion by Deshayes (1863). A specimen from Massawa, Eritrea (NHMUK) is likewise not a reliable record. Sowerby (1855) erroneously gave the locality of the species as Cuba. A much wider distribution was given by von Martens (1897 a), quoting records from the Ganges, Singapore, Borneo and Australia. Habitat and ecology. This species occupies a range of habitats, including salt marsh vegetation and mangroves on the fringes of salt pans and along creeks and rivers, in fully marine and brackish conditions. The snails climb up to 2 m on trunks, sometimes occurring in clusters of hundreds per tree, and descend to the ground to feed at low tide. Day (1974) recorded an occurrence up to 20 km from the mouth of the Morrumbene estuary (Mozambique), where salinity varied from 10 ppt at high water of spring tides (HWST) to 0.1 ppt at low water of spring tides (LWST). There have been several studies of vertical migration behaviour, but results are varied and sometimes contradictory, since observations have been made under different tidal conditions and sometimes for only short periods. Vannini et al. (2006) showed that behaviour depends upon the daily tidal cycle, spring-tide cycle, daynight cycle and tidal level within the mangrove forest, while Hodgson & Dickens (2012) reported seasonal differences in a temperate location. In Durban Bay, Brown (1971) observed aggregations of up to several hundred snails resting on single tree trunks, concentrated on the shadiest side, up to 1–2 m above the ground. They were most abundant in the Avicennia zone, but present also in the Bruguiera zone behind, and also found among marsh grasses and sedges. The animals did not graze on the trunks, but descended to the ground to feed during neap tides, climbing again at spring tides. There was only limited movement to other trees up to 5 m away during the course of a month. Observations were made in the dry season and behavioural patterns were not related to rainfall. Further details were provided by Cockcroft & Forbes (1981 a), working in the same area. They described a 14 -day tidal rhythm, with descent to the substrate during low tides being most frequent during neap tide periods, while at spring tides the snails tended to remain on trees throughout the tidal cycle. The snails descended more frequently in an upper intertidal area where inundation was less prolonged and began to climb up about 1 h before the flood tide reached the base of the tree. On the trunks they congregated at 0.5–0.7 m above the maximum tidal level. A series of detailed field studies was begun by Vannini et al. (2006) in the Avicennia belt between HWST and high water of neap tides (HWNT) in a mangrove forest in Kenya. The snails were never found on Lumnitzera or grass at higher levels and only rarely on Rhizophora trees on the seaward side. In the Avicennia zone there were usually 10–40 resting on each tree, but up to 350 could be found, close-packed together, just above the high-water mark up to 1 m above the ground. They rested at lower levels during neap tides and at the highest tidal levels were found only at the bases of trees. Most of the snails rested on tree trunks during high tide and descended to crawl and feed on the mud after the tide had receded, returning to the trunks before being reached by the advancing tide. This pattern was less pronounced at night, when many remained on the trunks throughout the low tide period. The migratory pattern was most evident during spring tides, whereas at neap tides the animals crawled on the mud regardless of tidal height or daylight. It also depended upon tidal level within the mangrove: at lower levels almost all snails migrated twice daily, but at upper levels, seldom reached by the tide, activity was more continuous and animals spent most of their time on the ground, even if covered with a few cm of water. An internal clock controls upward migration in advance of the rising tide, because migration begins when the tide is up to 100 m away (Vannini et al. 2008 c). The clock informs both the time and distance of migration; transplantation experiments showed that the behavioural rhythm was only reset to a new tidal level after 5–6 successive tides. All animals climbed to avoid submergence, but downward migration by individuals varied quite widely, suggesting that it is controlled not only by the internal clock, but also by direct cues from local conditions. The nature of these cues is unknown; neither odours, humidity nor vision would appear to be responsible (Vannini et al. 2008 a). Similarly, the cues that determine the height to which the snails climb to avoid the predicted tidal rise remain mysterious, despite experimental study (Vannini et al. 2008 b). Only animals larger than 9.5 mm showed migratory behaviour, while smaller individuals remained on the ground (Cockcroft & Forbes 1981 b; Vannini et al. 2006, 2008c). The southernmost limit of C. decollata is the Knysna estuary, where Hodgson & Dickens (2012) studied a population living at densities of up to 170 /m 2 among the rush Juncus. They noted that the species was first recorded in the estuary by Macnae (1963) and has since become abundant. As in mangrove habitats, the snails climbed the vegetation about 1 h before being contacted by the rising tide, although they did survive complete submersion when the Juncus was entirely covered by the highest spring tides. The snails were more active during the day and moved greatest distances in summer, while remaining inactive on the plants during the winter months. They did not return to the same resting site on Juncus plants after foraging, but did show homing to large wooden posts. This migratory behaviour clearly minimizes the danger of submergence and maximizes the time spent on the ground, but its adaptive value is debated. Submergence does not cause drowning, because animals survived for 14 days under aerated sea water (Cockcroft & Forbes 1981 a). Most authors have suggested that vertical migration is related to avoidance of predators that forage at high tide, such as the large mud crab Scylla serrata and predatory fish (Brown 1971; Cockcroft & Forbes 1981 b; Vannini et al. 2006, 2008a), which are more dangerous than the smaller eriphiid crab Epixanthus dentatus that snails encounter at low tide (Vannini et al. 2001). The majority of shells show the scars of unsuccessful attacks by shell-crushing predators (pers. obs.), so predators are expected to exert strong selection pressure. Migration could also be connected with physiological stress as suggested in C. anticipata by McGuiness (1994; but see remarks on that species above). Cockcroft & Forbes (1981 b) studied growth and mortality in Durban Bay. They estimated an age of 3 years at the modal size and 9 years at the asymptotic size. Copulatory behaviour was observed in September and October. Remarks. This is the sole species of Cerithidea in the western Indian Ocean, where it often occurs in high abundance. This is in marked contrast to most congeners, of which only C. moerchii sometimes approaches similar densities. The shell is quite variable, particularly in number of ribs, but is unlikely to be confused with any other, due to the diagnostic combination of fine spiral ridges, straight axial ribs and rounded whorls. In molecular phylogenetic analyses of COI, 16 S rRNA and 28 S rRNA its closest relationships were unresolved (Reid et al. 2013; Fig. 1), but in an analysis with more limited sampling but using 18 S rRNA, 28 S rRNA and COI, it appeared as sister to the large crown group of C. tonkiniana, C. moerchii, C. obtusa, C. anticipata, C. quoyii and C. dohrni (Reid et al. 2008). Genetic structure and shell variation along the East African coast have been studied by Madeira et al. (2012). The two were not closely correlated, but there were significant differences in shell shape among localities at a regional scale. It was suggested that large-scale genetic structure was consistent with a high dispersal potential., Published as part of Reid, David G., 2014, The genus Cerithidea Swainson, 1840 (Gastropoda: Potamididae) in the Indo-West Pacific region, pp. 1-65 in Zootaxa 3775 (1) on pages 43-47, DOI: 10.11646/zootaxa.3775.1.1, http://zenodo.org/record/285731, {"references":["Linnaeus, C. (1767) Systema Naturae per Regna tria Naturae. Ed. 12. Vol. 1. Part II. Stockholm, pp. 533 - 1327.","Gmelin, J. F. (1791) Caroli a Linne ... Systema Naturae per Regna Tria Naturae. Ed. 13. Vol. 1. Part 6. Leipzig, pp. 3021 - 3910.","Dillwyn, L. W. (1817) A Descriptive Catalogue of Recent Shells. Vol. 2. J. & A. Arch, London, pp. 581 - 1002.","Hanley, S. (1855) Ipsa Linnaei Conchylia. The Shells of Linnaeus, Determined from his Manuscripts and Collection. 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(Gastropoda: Prosobranchia: Cerithiidae). South African Journal of Zoology, 16, 5 - 9.","Vannini, M., Mrabu, E., Cannicci, S., Rorandelli, R. & Fratini, S. (2008 c). Rhythmic vertical migration of the gastropod Cerithidea decollata in a Kenyan mangrove forest. Marine Biology, 153, 1047 - 1053. http: // dx. doi. org / 10.1007 / s 00227 - 007 - 0877 - 8","Vannini, M., Coffa, C., Lori, E. & Fratini, S. (2008 a) Vertical migrations of the mangrove snail Cerithidea decollata (L.) (Potamididae) through a synodic month. Estuarine, Coastal and Shelf Science, 78, 644 - 648. http: // dx. doi. org / 10.1016 / j. ecss. 2008.02.010","Vannini, M., Lori, E. Coffa, C. & Fratini, S. (2008 b) Cerithidea decollata: a snail that can foresee the future? Animal Behaviour, 76, 983 - 992. http: // dx. doi. org / 10.1016 / j. anbehav. 2008.05.016","Cockcroft, V. G. & Forbes, A. T. (1981 b) Growth, mortality, and longevity of Cerithidea decollata (Linnaeus) (Gastropoda: Prosobranchia) from Bayhead mangroves, Durban Bay, South Africa. Veliger, 23, 300 - 308.","Vannini, M., Cannicci, S. & Fratini, S. (2001) Prey selection of Epixanthus dentatus (Crustacea: Brachyura: Eriphiidae) as determined by its prey remains. Journal of the Marine Biological Association of the United Kingdom, 81, 455 - 459. http: // dx. doi. org / 10.1017 / s 0025315401004088","McGuiness, K. A. (1994) The climbing behaviour of Cerithidea anticipata (Mollusca: Gastropoda): the roles of physical and biological factors. Australian Journal of Ecology, 19 (3), 283 - 289.","Madeira, C., Alves, M. J., Mesquita, N., Silva, S. E. & Paula, J. (2012) Tracing geographical patterns of population differentiation in a widespread mangrove gastropod: genetic and geometric morphometrics surveys along the eastern African coast. 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Cerithidea rhizophorarum A. Adams, 1855 (Figures 14 B, 15 A–I) Cerithidea rhizophorarum A. Adams, 1855: 85 –86 (Cagayan, province of Misamis, Mindanao [Cagayan de Oro, Mindanao, Philippines]; 3 syntypes NHMUK 1996432 / 4–6, Fig. 15 A, seen; Higo et al. 2001: 29, fig.). Sowerby, 1866: sp. 13, pl. 2, fig. 13 a, b (as rhizoperarum; as rhizoporarum in text of pl. 3 and index). Tapparone-Canefri, 1874: 142 –143, pl. 1, fig. 14 (as rhizosperarum). Hidalgo, 1904 –1905: 203–204. Dautzenberg & Fischer, 1905: 134 –135 (in part, includes C. moerchii; as rhizoporarum). Springsteen & Leobrera, 1986: 64, pl. 14, fig. 15. Higo et al., 2001: 29, fig. Reid et al., 2013: fig. 2 (map). Cerithium rhizophorarum — Sowerby, 1855: 886, pl. 186, fig. 273. Potamides (Cerithidea) rhizophorarum — Tryon, 1887: 162, pl. 33, fig. 67 (in part, includes C. quoyii; as rhizoporarum). von Martens, 1897 a: 189. Cerithium (Cerithidea) rhizophorarum — Kobelt, 1890 a: 53 –54, pl. 11, figs 3, 4 (Cecalupo 2005: pl. 31, fig. 13). Cerithidea (Cerithidea) rhizophorarum — Cecalupo, 2005: 316, pl. 31, fig. 3. Cecalupo, 2006: 136, 234 (in part, includes C. decollata, C. moerchii, C. weyersi). Cerithium (Cerithidea) freytagi Kobelt, 1893: 152 –154, pl. 29, figs 1, 2 (Philippines; types not found; Cecalupo 2005: pl. 31, fig. 3). Cerithidea rhizophorarum var. freytagi — Dautzenberg & Fischer, 1905: 135 (in part, includes C. moerchii; as freitagi). Taxonomic history. The name ‘ C. rhizophorarum ’ has been widely applied to C. moerchii in Japan, Korea, China and Vietnam (see synonymy of the latter). However, C. rhizophorarum was originally described from the Philippines and is morphologically distinct, as recently noted by Reid et al. (2013). Higo et al. (2001) noted that the status of the possible syntypes of C. rhizophorarum (NHMUK 1996432) was undetermined. This registration number includes nine specimens, three in each of three lots. In one of these lots there is an original label with the locality ‘Cagayan, Philippines’, and a label ‘ C. rhizophorarum A. Ad. ’ in the hand of A. Adams himself, so the three shells in this lot are identified as unequivocal syntypes (NHMUK 1996432 / 4–6) and the other six have no type status. Apart from refiguring of type specimens (Higo et al. 2001; Cecalupo 2005), only one original figure of this species has been published in more than a century (Springsteen & Leobrera 1986). The identity of Cerithidea similis, described by A. Adams (1855) from the Philippines, is unknown, but some aspects of the description match the present species (see Excluded and Doubtful Species, above). Diagnosis. Shell: periphery rounded, spire whorls only slightly rounded; elongated-pupoidal outline; aperture slightly thickened and flared, sinuous apertural margin; 18–32 rounded axial ribs on penultimate whorl, ribs usually weak or absent after ventrolateral varix; ventrolateral varix a wide rib at 170–270 °; 0–7 varices on spire; 5– 6 spiral cords above periphery; brown, paler at shoulder. Philippines. Material examined. 18 lots. Shell (Fig. 15 A–I): H = 18.0– 36.1 mm. Shape elongated-pupoidal (H/B = 2.14–2.66; SH = 3.0– 3.52); decollate, 6–7 whorls remaining; spire whorls slightly rounded, suture sometimes indistinct; spire profile convex, final whorl constricted so adults have pupoidal outline; 0–7 thick pale varices on spire; periphery rounded; moderate thickness. Adult lip slightly thickened and flared; apertural margin sinuous in side view; weak anterior projection adjacent to canal. Sculpture on spire of straight to slightly opisthocline axial ribs, becoming slightly curved (opisthocyrt) on later whorls, ribs low, rounded, interspaces equal to width of ribs, 18–32 ribs on penultimate whorl, ribs weak or absent after ventrolateral varix; spiral sculpture of 5 strong, flattened, primary cords on spire whorls, of which anterior cord is usually divided on later whorls to give 6 cords above periphery, posterior cord adjacent to suture is strongest and bears small nodules (corresponding to, or up to twice as numerous as axial ribs); spiral cords cross relatively weaker axial ribs; base with 8–10 flattened cords or ridges. Ventrolateral varix a wide rib at 170–270 °; 0–7 wide varices on spire. Surface with fine spiral microstriae on periostracum; surface dull when worn. Colour: white, fawn or pale brown, often with paler zone across 2–3 posterior cords; spiral grooves darker brown, varices white; aperture pale brown. Animal: unknown. Range (Fig. 14 B): Philippines: Subic Bay, Luzon (USNM 593859); Mactan I., Cebu (USNM 862804); Panabutan Bay, Mindanao (USNM 862797); Cagayan de Oro, Mindanao (NHMUK 1996432); Nabas, Panay (S. Tracey); Palawan (Springsteen & Leobrera 1986). According to Springsteen & Leobrera (1986: 64) this species occurs “throughout the Philippines ”. Sowerby (1866) gave ‘Borneo and the Philippines’ as the distribution, but occurrence in Borneo has not been verified and is considered unlikely in view of the many reliable records of other Cerithidea species from the island. Kobelt’s (1890 a) record from northern Australia is incorrect. Habitat and ecology. On sandy mud near the estuary of a small drainage stream from a mangrove area at Nabas, Panay (S. Tracey, pers. comm.). Remarks. Until very recently (Reid et al. 2013), ‘ C. rhizophorarum ’ has been well known as an oriental species found from northern Honshu, throughout the coast of China and to Vietnam. However, the type locality of A. Adams’ species is Mindanao in the Philippines and it would be unusual for a predominantly warm-temperate oriental species to extend its range across the South China Sea to the tropical Philippines. Close comparison of shells revealed differences, which persuaded Reid et al. (2013) to treat the form from the Philippines as a distinct species, C. rhizophorarum s. s., while the mainland species was identified as C. moerchii. The differences are subtle. Cerithidea rhizophorarum has a pupoidal outline (i.e. convex spire profile), the whorls are nearly flat, the axial ribs are fine and numerous, and the colour is pale with broad light brown zones (without dark, narrow bands) (Fig. 15 A–I). Cerithidea moerchii has a straight spire, more rounded whorls, often shows darker coloration and sometimes has rugose axial ribs (Fig. 15 J–AA). Significantly, the shells of C. moerchii from the Ryukyu Islands and Taiwan, which from their geographical proximity could be expected to be most similar if all were conspecific, are in fact the most distinct from C. rhizophorarum in the Philippines (the shells from Ryukyus and Taiwan are small and rugose, Fig. 15 U–AA; see Remarks on C. moerchii). So far, no molecular samples have been available to test this hypothesis, but it is predicted that C. rhizophorarum and C. moerchii will prove to be distinct sister species. The intertidal shells of the Philippines are rather well known, so it is surprising that the presence there of C. rhizophorarum has been mentioned in the literature only once in the past century, when Springsteen & Leobrera (1986: 64) reported that it occurred “throughout the Philippines ”. The species was not included in the family account by Lozouet (2008) in the comprehensive Philippine Marine Mollusks and was not found by Lozouet & Plaziat (2008) in Bohol. There are, however, 15 lots from the Philippines in USNM, mostly collected by P. Bartsch in the early twentieth century. Two of these lots consist of> 100 specimens and one of these was from Batangas Market, Luzon (USNM 243643), suggesting that the snails had been collected for food. A few specimens were found in Panay in 1979 by S. Tracey (pers. comm.). The conservation status of this species could be a matter of concern., Published as part of Reid, David G., 2014, The genus Cerithidea Swainson, 1840 (Gastropoda: Potamididae) in the Indo-West Pacific region, pp. 1-65 in Zootaxa 3775 (1) on pages 50-52, DOI: 10.11646/zootaxa.3775.1.1, http://zenodo.org/record/285731, {"references":["Adams, A. (1855) A monograph of Cerithidea, a genus of Mollusca, with descriptions of several new species, from the collection of High Cuming, Esq,: to which are added, descriptions of two new species of Colina, and one of Donax. Proceedings of the Zoological Society of London, 1854, 83 - 87. http: // dx. doi. org / 10.1111 / j. 1469 - 7998.1854. tb 07236. x","Higo, S., Callomon, P. & Goto, Y. (2001) Catalogue and Bibliography of the Marine Shell-Bearing Mollusca of Japan. Gastropoda. Bivalvia. Polyplacophora. Scaphopoda. Type Figures. Elle Scientific Publications, Osaka, 208 pp.","Sowerby, G. B. II (1866) Monograph of the genus Cerithidea. In: Reeve, L. A. (Ed.), Conchologia Iconica. Vol. 15. L. A. Reeve, London, pls 1 - 4.","Tapparone-Canefri, C. (1874) Zoologia del viaggio intorno al globo della R. Fregata Magenta. Malacologia. Memorie della Reale Accademia delle Scienze di Torino, Ser. 2, 28, 109 - 265.","Hidalgo, J. G. (1904 - 1905) Catalogo de los moluscos testaceos de las Islas Filipinas, Jolo y Marianas. I. - Moluscos marinos. Gaceta de Madrid, Madrid, xvi + 408 pp.","Dautzenberg, P. & Fischer, H. (1905) Liste des mollusques recoltes par M. le Capitaine de Fregate Blaise au Tonkin, et description d'especes nouvelles. Journal de Conchyliologie, 53, 85 - 234.","Springsteen, F. J. & Leobrera, F. M. (1986) Shells of the Philippines. Carfel Seashell Museum, Manila, 377 pp.","Reid, D. G., Claremont, M., Smith, L., Shamoto, M., Glaubrecht, M. & Ozawa, T. (2013) Mosaics in the mangroves: allopatric diversification of tree-climbing mudwhelks (Gastropoda: Potamididae: Cerithidea) in the Indo-West Pacific. Biological Journal of the Linnean Society, 110, 564 - 580.","Sowerby, G. B. II (1855) Monograph of the genus Cerithium, Adanson. In: Sowerby, G. B. II. (Ed.), Thesaurus Conchyliorum, or Figures and Descriptions of Recent Shells. Vol. 2. part 16. Sowerby, London, pp. 847 - 899.","Tryon, G. W. (1887) Manual of Conchology. Vol. 9. Tryon, Philadelphia, 488 pp.","Martens, E. von (1897 a) Suss- und Brackwasser-Mollusken des Indischen Archipels. In: Weber, M. (Ed.), Zoologische Ergebnisse einer Reise in Niederlandisch Ost-Indien, Vol. 4. E. J. Brill, Leiden, pp. 1 - 331.","Kobelt, W. (1890 b) Eine neue Cerithidea. Nachrichtsblatt der deutschen Malakozoologischen Gesellschaft, 1890, 60 - 61.","Cecalupo, A. (2005) Elenco della famiglia Cerithiidae Ferussac, 1822 (Prosobranchia). Nomenclatura delle specie conosciute o poco note e relativa revisione sistematica. Fossili e Attuali. Vol. I. Quaderni della Civica Stazione Idrobiologica di Milano, 26, 1 - 366.","Cecalupo, A. (2006) Elenco della famiglia Cerithiidae Ferussac, 1822 (Prosobranchia). Nomenclatura delle specie conosciute o poco note e relativa revisione sistematica. Attuali. Vol. II. Quaderni della Civica Stazione Idrobiologica di Milano, 27, 1 - 373.","Lozouet, P. & Plaziat, J. - C. (2008) Mangrove Environments and Molluscs. Abatan River, Bohol and Panglao Islands, Central Philippines. ConchBooks, Hackenheim, 160 pp."]}

Reid, David G. (2014): The genus Cerithidea Swainson, 1840 (Gastropoda: Potamididae) in the Indo-West Pacific region. Zootaxa 3775 (1): 1-65, DOI: http://dx.doi.org/10.11646/zootaxa.3775.1.1

Cerithidea anticipata Iredale, 1929 (Figures 2 C, 10, 11) Cerithium obtusum var. b Lamarck, 1822: 71 (MHNG INVE 51622 (formerly MHNG 1097 / 24), photograph seen; not Cerithium obtusum Lamarck, 1822). Cerithium obtusum — Kiener, 1841 –1842: 95–96, pl. 29, fig. 2 (in part, includes C. obtusa; not Lamarck, 1822). Cerithidea obtusa — Cernohorsky, 1972: 60, pl. 13, fig. 1 (in part, includes C. obtusa; not Lamarck, 1822). Healy, 1983: 57–75, figs 62, 95– 98 (euspermatozoa; not Lamarck, 1822). Cerithidea (Cerithidea) obtusa — Wilson, 1993: 133, pl. 15, fig. 15 (not Lamarck, 1822). Cerithium kienerii Hombron & Jacquinot, 1848: pl. 23, figs 4, 5 (living animal) (Rafles-Bay [Raffles Bay, Cobourg Peninsula, Northern Territory, Australia]; holotype MNHN 25692, Fig. 10 A; as C. kennerii; not Cantraine, 1835). Rousseau, 1854: 96 –97 (as kieneri). Sowerby, 1855: 886, pl. 186, fig. 272 (as kieneri). von Martens, 1897 a: 188 (as kennerii). Cerithidea kieneri —H. Adams & A. Adams, 1854: 293. Sowerby, 1866: sp. 6, pl. 1, fig. 6. Hedley, 1909: 357. Allan, 1950: 87 – 88. Potamides (Cerithidea) obtusa var. kieneri — Tryon, 1887: 161, pl. 32, fig. 53, pl. 33, fig. 61. Cerithium (Cerithidea) kieneri — Kobelt, 1893: 159–160, pl. 30, fig. 1. Cerithidea multicostata Schepman, 1919: 192, pl. 8, fig. 3, 3a (Merauke, New Guinea [East Papua, Indonesia]; syntypes ZMA 136547). Van der Bijl et al., 2010: 72, fig. 178. Cerithidea anticipata Iredale, 1929: 278 (new name for Cerithium kieneri Hombron & Jacquinot, 1848, not Cantraine, 1835). Houbrick, 1986: 284, 285, figs 5, 6, fig. 15 (radula). Cecalupo, 2005: 316, pl. 31, fig. 21. Cecalupo, 2006: 27, 176, pl. 57, fig. 13 a–c (in part, includes C. balteata, Cerithideopsilla conica, Cerithideopsis largillierti). Reid et al., 2008: 680 –699, figs 1, 2 (phylogeny). Jarrett, 2011: 40, fig. 95. Willan, 2013: 77, fig. 8. Reid et al., 2013: figs 1 (shell, phylogeny), 2 (map). Cerithidea (Cerithidea) anticipata — Wilson, 1993: 132 –133, pl. 15, fig. 16. Cerithidea reidi — Cecalupo, 2006: 85, 210, 233– 234 (not Houbrick, 1986; in part, includes C. reidi, C. dohrni). Taxonomic history. This species was first noticed as ‘var. b’ of Cerithium obtusum by Lamarck (1822; see Taxonomic History of that species) and subsequently figured under the same name by Kiener (1841 –1842). This was presumably the reason why Hombron & Jacquinot (1848) named the species Cerithium kienerii, which they distinguished from Cerithium obtusum by the different body coloration. Nevertheless, the name C. obtusa has occasionally been used for this Australasian species in more recent literature (Cernohorsky 1972; Healy, 1983; Wilson 1993). The authorship and spelling of Cerithium kienerii require justification. As demonstrated by Clark & Crosnier (2000), the Atlas of the Voyage au Pole Sud et dans l’Océanie sur les corvettes l’Astrolabe et la Zélée was published in parts (pl. 23 in 1848) by Hombron & Jacquinot, sometimes before the accompanying text. The written descriptions of molluscs were provided by Rousseau and published in 1854. According to the ICZN Code (1999: Art.12.2.7) an illustration with a binominal name constitutes a valid indication, so in this case the new names in the Atlas are available and take priority over those in the text. In its original form in the caption to plate 23 the specific name was spelled kennerii, but this appeared as kieneri in Rousseau’s text. This is taken as evidence of an incorrect original spelling (ICZN 1999: Art. 32.5.1), even though the correction was made in a subsequent volume of the original work, since the subsequent text was prepared under the supervision of the original authors. The original genitive ending –ii is not to be changed, however (ICZN 1999: Art. 33.4). An alternative interpretation is that the correction to kieneri was an incorrect subsequent spelling that cannot be used as a substitute name (ICZN 1999: Art. 33.3); this would have the undesirable consequence that kennerii would become the valid name for this species. The original form of the name, kennerii, has in fact been used only once, by von Martens (1897 a). Several early authors (H. Adams & A. Adams 1854; Sowerby 1855, 1866; Tryon 1887) evidently recognized that kennerii was an error, and attributed the corrected version kieneri to Hombron & Jacquinot, and this adds to the argument that the latter (but with the – ii ending) is the correct original spelling (ICZN 1999: Art. 33.3.1). During the twentieth century the corrected name was used at least twice, by Hedley (1909) and Allan (1950). At present this species is familiar as C. anticipata, a replacement name provided by Iredale (1929) for the preoccupied Cerithium kienerii. In fact this replacement may not have been necessary, because an older available name exists: C. multicostata Schepman, 1919. This was based on the form with numerous axial ribs from southern New Guinea. Comparison of shells suggests that this form is conspecific with those from Australia, but genetic distances within Australia indicate that more than one species could exist within a complex (see Remarks). Until this is clarified by new data, it seems premature to change the familiar name C. anticipata. Diagnosis. Shell: periphery rounded or weakly angled; aperture flared, anterior canal and apertural projection well developed; 14–40 axial ribs on penultimate whorl; ventrolateral varix a slightly enlarged rib at 200–260 °; 5 spiral cords on spire, 4–10 above periphery on last whorl; brown with darker spiral cords. Northern Australia and New Guinea. COI GenBank AM 932760 – AM 932762. Material examined. 57 lots. Shell (Fig. 10 A–P): H = 24.0– 50.3 mm. Shape elongated conical (H/B = 2.01–2.62, SH = 2.83–3.54); decollate, 7–10 whorls remaining; spire whorls rounded, suture distinct; spire profile straight to slightly convex; periphery rounded or weakly angled; moderately solid. Adult lip flared, moderately thickened; apertural margin planar in side view; strong anterior projection adjacent to deep notch of anterior canal. Sculpture on spire of straight to slightly curved (opisthocyrt) axial ribs, often bifurcating posteriorly (adapically) to give a row of granules at suture (occasionally bifurcation occurs lower on whorl to increase number of axial ribs; Fig. 10 J, N–P); ribs prominent, narrowly rounded, interspaces 1.5–2 times width of ribs (occasionally equal), 14–40 ribs on penultimate whorl, remaining strong but becoming more distant after ventrolateral varix, weak on base; spire whorls with 5 narrow spiral cords, often forming nodules where they cross axial ribs, usually increasing by interpolation of narrower threads to give 4–10 spiral cords and threads above periphery on final whorl; base with 7–11 narrow ridges, peripheral one is a cord of same size as primary cords above, but others on base slightly narrower. Ventrolateral varix a slightly enlarged rib at 200–260 °. Surface with fine spiral microstriae on periostracum. Colour: pinkish brown to fawn, spiral ribs usually darker brown (especially on anterior half of whorls, giving appearance of broad brown band), axial ribs paler; aperture pale brown, spiral lines showing through. Animal (Fig. 2 C): Head and base of tentacles pinkish grey with cream spots; anterior half of snout blackish with scattered yellow spots; tentacles cream, grey towards tip; sides of foot dark grey speckled with grey, brown and white, small yellow spots at margin; sole of foot grey with pinkish border; mantle blue grey. Range (Fig. 11): Queensland (Qld) and Northern Territory (NT) of Australia, New Guinea. Records: Australia: Prince Regent River Reserve, Western Australia (Houbrick 1986: fig. 5); Forrest River Mission, 50 miles from Wyndham, NT (AM C.085156); Buffalo Creek, Darwin, NT (NHMUK 20130241; USNM 828819); Bing Bong Station, W Gulf of Carpentaria, NT (AM C. 412929); Allen I., S Wellesley Is, S Gulf of Carpentaria, Qld (AM C.015839); Moa I., Torres Strait, Qld (NHMUK 20130244); Cockle Bay, Magnetic I., Qld (NHMUK; USNM 828828); Lota Creek, Wynnum, Moreton Bay, Qld (AM C. 138069); Brunswick R., New South Wales (D. Riek, pers. comm.). Indonesia: Kupang, West Timor (ZMB); Biak, Soepiori I., Schouten Is, East Papua (ANSP 207534); Sulawati I., East Papua (AM); Kokenau, Mimika R., East Papua (AM C. 458258); Merauke, East Papua (AM C. 121289). Papua New Guinea: Collingwood Bay (AM C. 18891). Sowerby (1855, 1866) erroneously gave the locality as ‘ Philippines and Borneo’ (as Cerithium kieneri). Habitat and ecology. This species is typically found in the Ceriops zone of mangrove forests, often among shrubs on the fringes of bare salt pans, at heights of up to 1 m on trunks, and sometimes on firm mud beneath the trees. It is less common on trunks in the Rhizophora and Bruguiera zones, or in the landward fringe, where it may reach 3 m up the trees. In a detailed study at Darwin, McGuiness (1994) found C. anticipata to be most abundant in the mangrove zone dominated by Ceriops, often in and around clearings with dead trees (caused by cyclone damage). The animals rested in shady sites on trees, attached by dried mucus with only a small part of the outer lip in contact with the trunk; they climbed higher, and were less active, during neap tides that did not flood the forest than during spring tides. Snails that were tethered to prevent climbing died, apparently from physiological stress (but these were tethered during neap tides in an open clearing where soil temperature reached 45 °C, so these results may not be typical). It was suggested that, consequently, the major reason for the climbing behaviour was avoidance of physiological stress, not escape from subtidal predators that enter the mangrove during spring tides. Nevertheless, predation was evident, because many snails showed signs of shell damage. Studies of C. decollata have suggested that predator avoidance has led to the evolution of climbing behaviour (Cockcroft & Forbes 1981 a; Vannini et al. 2006). McGuiness (1994) argued that the Darwin site was at a higher tidal level than the locations of studies of C. decollata, so that physiological stress was more significant. In the vicinity of Darwin, Willan (2013: 77) reported that C. anticipata can be found in both mangrove and salt marsh environments, and occurs in “extremely high densities” among the foliage of the canopy, where it is quiescent during the drier months. He noted that it is rarely found on the ground, mainly foraging at low levels on the trunks of various mangrove tree species, when conditions are moist. However, McGuiness (1994) implied that activity takes place mainly at night, so daytime observations may not be representative. Remarks. The shell shows considerable regional variation. Specimens from Merauke, S New Guinea, and one from the nearby Torres Strait (Fig. 10 J, N–P) show increased numbers of fine axial ribs on the spire (these were named C. multicostata Schepman, 1919). Shells from elsewhere in New Guinea are slender and delicate, but again have fine and sometimes numerous axial ribs (Fig. 10 E, F). Shells from E Queensland are large, with coarse sculpture (Fig. 10 H, I, K–M). Those from the Gulf of Carpentaria are thick and pale. In the Northern Territory the whorls are slightly compressed vertically to produce a broader shell (Fig. 10 A–D). COI sequence data reveal a deep division between two specimens from Darwin and one from the Torres Strait (uncorrected pairwise distance = 0.072) and phylogenetic analysis does not give significant support to the monophyly of all three (Reid et al. 2013; Fig. 1). However, molecular sampling is so far too limited to determine whether this division corresponds in any way with the observed pattern of morphological variation. The distribution appears to be continuous around the northern coast of Australia, so does not suggest any geographical separation between eastern and western populations. Nevertheless, a pattern of eastern and western genetic differentiation (both intra- and inter-specific) across northern Australia is common in marine organisms and has been explained by isolation during glacial low sea-level stands (Reid et al. 2006, 2010). Further investigation is required of genetic structure and possible speciation in C. anticipata. The sister species of C. anticipata is C. reidi (Reid et al. 2013; Fig. 1). Morphologically, C. anticipata is distinguished from C. reidi by smaller size, darker coloration and narrower ribs and cords (see Remarks on C. reidi). However, the genetic distance between them is similar to that between the divergent sequences of C. anticipata (uncorrected pairwise distance for COI = 0.076, cf 0.072 within C. anticipata). Together, all form a wellsupported clade (Fig. 1). Houbrick (1986) reported that the ranges of these two species just overlap in Admiralty Gulf in northern Western Australia. He noted that, in the region of overlap, specimens of C. anticipata “are larger than specimens from northern Australia and Queensland and are more similar to C. reidi in shell sculpture. These larger C. anticipata, while resembling C. reidi, never have a purple aperture and do not attain the large size of C. reidi. These two species are probably very closely related, as their radulae and shell sculptures are morphologically similar even though there is great size disparity” (Houbrick 1986: 285). He illustrated a shell that appears to be C. anticipata, from Prince Regent River (Houbrick 1986: fig. 5), which is just within the geographical range of C. reidi. Nevertheless, it is possible that some of the shells of ‘ C. anticipata ’ he observed from Admiralty Gulf were actually small specimens of C. reidi (see Remarks on C. reidi) and their respective ranges in this area remain to be confirmed. Detailed genetic study is required to test whether these are distinct species, or whether hybridization occurs, leading to intergradation where they overlap. In eastern New Guinea the range of C. anticipata overlaps that of C. balteata and shells of these two species can be very similar. Cerithidea anticipata can be distinguished by its stronger spiral sculpture on the spire whorls (Fig. 5 Q–U, cf. Fig. 10 E–G; see Remarks on C. balteata). Aboriginal people in Australia do not use C. anticipata as food (Willan 2013)., Published as part of Reid, David G., 2014, The genus Cerithidea Swainson, 1840 (Gastropoda: Potamididae) in the Indo-West Pacific region, pp. 1-65 in Zootaxa 3775 (1) on pages 32-36, DOI: 10.11646/zootaxa.3775.1.1, http://zenodo.org/record/285731, {"references":["Iredale, T. (1929) Queensland molluscan notes, no. 1. Memoirs of the Queensland Museum, 9, 261 - 297.","Lamarck, J. B. P. A. de (1822) Histoire Naturelle de Animaux sans Vertebres. Vols 6 (2), 7. 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(1986) Cerithidea reidi, spec. nov., from Western Australia. Veliger, 28, 280 - 286.","Cecalupo, A. (2005) Elenco della famiglia Cerithiidae Ferussac, 1822 (Prosobranchia). Nomenclatura delle specie conosciute o poco note e relativa revisione sistematica. Fossili e Attuali. Vol. I. Quaderni della Civica Stazione Idrobiologica di Milano, 26, 1 - 366.","Cecalupo, A. (2006) Elenco della famiglia Cerithiidae Ferussac, 1822 (Prosobranchia). Nomenclatura delle specie conosciute o poco note e relativa revisione sistematica. Attuali. Vol. II. Quaderni della Civica Stazione Idrobiologica di Milano, 27, 1 - 373.","Reid, D. G., Dyal, P., Lozouet, P., Glaubrecht, M. & Williams, S. T. (2008) Mudwhelks and mangroves: the evolutionary history of an ecological association (Gastropoda: Potamididae). Molecular Phylogenetics and Evolution, 47, 680 - 699. http: // dx. doi. org / 10.1016 / j. ympev. 2008.01.003","Jarrett, A. G. (2011) Marine Shells of the Whitsunday Coast & Islands. Carole Green Publishing, Cambridge, v + 268 pp.","Willan, R. C. (2013) A key to the potamidid snails (longbums, mudcreepers and treecreepers) of northern Australia. Northern Territory Naturalist, 24, 68 - 80.","Reid, D. G., Claremont, M., Smith, L., Shamoto, M., Glaubrecht, M. & Ozawa, T. (2013) Mosaics in the mangroves: allopatric diversification of tree-climbing mudwhelks (Gastropoda: Potamididae: Cerithidea) in the Indo-West Pacific. Biological Journal of the Linnean Society, 110, 564 - 580.","Clark, P. F. & Crosnier, A. (2000) The zoology of the Voyage au pole Sud et dans l'Oceanie sur les corvettes l'Astrolabe et la Zelee execute par ordre du roi pendant les annees 1837 - 1838 - 1839 - 1840 sous le commandement de M. Dumont-d'Urville (1842 - 1854): titles, volumes, plates, text, contents, proposed dates and anecdotal history of the publication. Archives of Natural History, 27, 407 - 435. http: // dx. doi. org / 10.3366 / anh. 2000.27.3.407","McGuiness, K. A. (1994) The climbing behaviour of Cerithidea anticipata (Mollusca: Gastropoda): the roles of physical and biological factors. Australian Journal of Ecology, 19 (3), 283 - 289.","Cockcroft, V. G. & Forbes, A. T. (1981 a) Tidal activity rhythms in the mangrove snail Cerithidea decollata (Linn.) (Gastropoda: Prosobranchia: Cerithiidae). South African Journal of Zoology, 16, 5 - 9.","Vannini, M., Rorandelli, R., Lahteenoja, O., Mrabu, E. & Fratini, S. (2006) Tree-climbing behaviour of Cerithidea decollata, a western Indian Ocean mangrove gastropod (Mollusca: Potamididae). Journal of the Marine Biological Association of the United Kingdom, 86, 1429 - 1436. http: // dx. doi. org / 10.1017 / s 0025315406014470","Reid, D. G., Lal, K., Mackenzie-Dodds, J., Kaligis, F., Littlewood, D. T. J. & Williams, S. T. (2006) Comparative phylogeography and species boundaries in Echinolittorina snails in the central Indo-West Pacific. Journal of Biogeography, 33, 990 - 1006. http: // dx. doi. org / 10.1111 / j. 1365 - 2699.2006.01469. x","Reid, D. G., Dyal, P. & Williams, S. T. (2010) Global diversification of mangrove fauna: a molecular phylogeny of Littoraria (Gastropoda: Littorinidae). Molecular Phylogenetics and Evolution, 55, 185 - 201. http: // dx. doi. org / 10.1016 / j. ympev. 2009.09.036"]}

Cerithidea obtusa (Lamarck, 1822) (Figures 2 B, 8, 9) Cerithium obtusum Lamarck, 1822: 71 (“les mers de Timor”, probably in error, see Range below; lectotype MHNG INVE 51623 (formerly 1097 / 25), here designated, 43.8 mm, Fig. 8 A; in part, includes C. anticipata). Kiener, 1841 –1842: 95–96, pl. 29, fig. 1 (in part, includes C. anticipata). Hombron & Jacquinot, 1848: pl. 23, fig. 3 (living animal). Souleyet, 1852: 600, Atlas pl. 39, figs 1, 2 (living animal). Rousseau, 1854: 96. Sowerby, 1855: 885, p. 186, fig. 271. Hanley, 1856: 215, suppl. pl. 4, fig. 8. Strombus obtusus — Wood, 1828: 13, pl. 4, fig. 8. Hanley, 1856: 215, suppl. pl. 4, fig. 8. Cerithidea obtusa — H. Adams & A. Adams, 1854: 292–293, pl. 31, fig. 2 b. Adams, 1855: 83. Sowerby, 1866: sp. 4, pl. 1, fig. 4 a, b. Morlet, 1889: 144. Van Benthem Jutting, 1956: 433 –435, fig. 106. Habe & Kosuge, 1966: 25, pl. 7, fig. 23. Subba Rao & Mookherjee, 1975: 170. Subba Rao, Dey & Barua, 1983: 51, pl. 2, fig. 2. Houbrick, 1986: 284, figs 7, 8, figs 10 and 14 (radula). Subba Rao, Dey & Barua, 1992: 177, pl. 5, figs 3, 6. Poutiers, 1998: 450, fig. Dharma, 2005: 92, pl. 21, fig. 8. Thach, 2005: 49, pl. 10, fig. 4. Dey, 2006: 35 –36, figs 38–40. Ahmed & Mahmoud, 2007: 9, 95, figs pp. 9, 95 (in part, includes C. weyersi). Reid et al., 2008: 680 –699, figs 1, 2 (phylogeny), 2 H. Hamli et al., 2013: 412 –418, fig. 2 a. Reid et al., 2013: figs 1 (shell, phylogeny), 2 (map). Potamides (Cerithidea) obtusus —von Martens, 1880: 282. von Martens, 1887: 168. Tryon, 1887: 161, pl. 33, figs 59–60 (as obtusa; in part, includes C. quoyii, C. anticipata). von Martens, 1897 a: 186 –187; pl. 9, fig. 22; pl. 10, fig. 5 (head). von Martens, 1897 b: 266. Fischer & Dautzenberg, 1904: 416. Oostingh, 1923: 76 –77, fig. 12. Dautzenberg, 1929: 489. Cerithium (Cerithidea) obtusum — Kobelt, 1890 a: 42 –43, pl. 9, figs 3–5 (Cecalupo 2005: pl. 31, fig. 10). Potamides obtusum — Prashad, 1921: 495. Cerithidea (Cerithidea) obtusa — Brandt, 1974: 192 –193, pl. 14, fig. 52. Houbrick, 1984: 15, 16, fig. 5 C. Subba Rao, Surya Rao & Maitra, 1991: 44. Subba Rao, Dey & Barua, 1995: 45, 58. Glaubrecht, 1997: 299, pl. 12, fig. 6. Subba Rao, 2003: 136, pl. 22, figs 5, 6. Cecalupo, 2005: 316, pl. 31, fig. 10. Cecalupo, 2006: 116, 223, pl. 57, fig. 12 a–c. Ramakrishna et al., 2007: 47. Cerithium decollatum —Sowerby, 1834: pl. 214, fig. 2 (not Linnaeus, 1767). Reeve, 1842: 178, pl. 227, fig. 2 (not Linnaeus, 1767). Melania lineolata Gray in Griffith & Pidgeon, 1833: pl. 14, fig. 4 (no locality; holotype NHMUK 20130227, seen; not Wood, 1828). Cerithidea lineolata — Swainson, 1840: 342. Cerithium truncatum ‘Lam.’ Griffith & Pidgeon, 1834: 596, 598 (refers to pl. 14, fig. 4; as Cerethium p. 598; no locality; holotype NHMUK 20130227, seen; not Gray in Griffith & Pidgeon, 1833: pl. 13, fig. 1). Cerithidea ornata — Li, 2008: 82, fig. (not Sowerby, 1855). Taxonomic history. This species has been recognized under its original specific name by almost all authors, although there was some early misidentification as C. decollata (e.g. Sowerby 1834; Reeve 1842). The original concept of the species was broader than today; Lamarck’s (1822) description of his Cerithium obtusum included a ‘var. b’, which referred to C. anticipata. This was followed by Kiener (1841 –1842) and Tryon (1887) also had a broad concept of the species. As recorded by Lamarck’s daughter Rosalie (in an annotated copy of his Histoire Naturelle de Animaux sans Vertèbres), his collection contained a single specimen of the typical form of Cerithium obtusum and two of var. b (the latter, registered as MHNG INVE 51622, formerly MHNG 1097 / 24, are C. anticipata and have no type status; ICZN 1999: Art. 72.4.1). There are, however, currently two specimens in the lot of the typical form, one an adult (H = 43.8 mm; Fig. 8 A) and the other a juvenile (H = 37.8 mm) (Y. Finet, pers. comm.). Lamarck (1822) gave “ 19 lignes” (= 42.7 mm) for the typical form, so the larger shell corresponds most closely with his description. It may be the holotype but, to avoid any potential for future ambiguity, it is here designated lectotype. This is the type species of the genus Cerithidea, but the designation refers to its synonym, Melania lineolata, and the case is a complicated one. The molluscan taxa introduced in volume 12 of Griffith’s edition of Cuvier’s Animal Kingdom have been discussed in detail by Petit & Coan (2008), including their authorship and dates of publication. Briefly, some of the molluscan plates (including plates 13 and 14) were newly drawn for the molluscan volume (Griffith & Pidgeon 1833 –1834), from specimens in NHMUK provided by Gray. While Gray may also have supplied the names used in the legends (i.e. text appearing on the plates themselves), the index (an alphabetical list of species figured in the plates) was prepared by someone else and contains “many errors and misattributions of species” (Petit & Coan 2008: 222). In their commentary on the availability of names, Petit & Coan (2008) adhered strictly to the published evidence. Thus in the legend of plate 14 the shell illustrated in figure 4 (here shown to be C. obtusa, see below) is labelled Melania lineolata, which they interpreted as an available but invalid name (a junior homonym of Melania lineolata Wood, 1828) attributed to Gray and published in 1833. Previous authors, including Swainson (1840), Bequaert (1942 a, b) and Houbrick (1984), have also reached the same conclusion. Since Wood’s species was illustrated and correctly labelled on the preceding plate 13, and because the genera Melania and Cerithium (in a broad sense, to include Cerithiidae and Potamididae) were clearly diagnosed in the main text and would not have been confounded by the experienced Gray, it seems more likely that the legend of plate 14, figure 4, was simply an error. This error was corrected in the index of the volume, published the following year (Griffith & Pidgeon 1834). On page 596 the caption to plate 14, figure 4, is given as “ Cerithium truncatum, Lam. ” and this is repeated on page 598 (without the attribution); Bequaert (1942 b) and Petit & Coan (2008) have interpreted this action as a replacement name for the preoccupied Melania lineolata. Again, without evidence of intent by the original authors, this seems questionable, because the name was attributed to Lamarck (who introduced no such name); more plausibly, this was another error, for Lamarck’s Cerithium obtusum, a name of similar descriptive meaning. To compound the confusion, the name Cerithium truncatum had been used for plate 13, figure 1, and was subsequently renamed “ Cerithium laeve, Gray ” in the index (this is the species now known as Campanile symbolicum Iredale, 1917). Both Iredale (1917: 326) and Houbrick (1981: 283) described the use of Cerithium truncatum in this latter context as “a careless slip” and did not accept it as an available name. However, Petit & Coan (2008) did accept it as available and therefore a senior homonym of the replacement name for the shell of plate 14, figure 4. Regardless of this speculation about the original intent of Gray and likelihood of errors in the publication of Griffith & Pidgeon, it is recommended that the strict interpretation of Petit & Coan (2008) should be followed (see Synonymy above). This has no effect on the valid name of the present species, since both the synonyms are junior primary homonyms. It does, however, have an implication for the concept of the genus Cerithidea because, in introducing the genus, Swainson (1840) gave “ C. lineolata. Griff. Cuv. 14. f. 4 ” (i.e. Melania lineolata Gray in Griffith & Pidgeon, 1833) as one of only two originally included species (the other being a fossil), which was subsequently designated type species by Makiyama (1936). For stability, it is therefore important to accept this name as available. If considered unavailable from Griffith & Pidgeon (1833), the name could still be accepted as available as Cerithidea lineolata Swainson, 1840, because Swainson’s reference to the published figure is a valid indication of the name (ICZN 1999: Art. 12.2.7). The original figure of Melania lineolata Gray in Griffith & Pidgeon, 1833 shows a rather elongate spire and the apertural lip is neither extremely flared nor strongly thickened. For these reasons Reid (quoted in Petit & Coan 2008) suggested it might represent C. reidi, rather than C. obtusa as previously interpreted (e.g. Houbrick 1984). The original shell has since been discovered (NHMUK 20130227), proving that the figure was poorly drawn from an oblique angle and that it is indeed a specimen of C. obtusa. Diagnosis. Shell: large, solid, broad, periphery rounded; aperture strongly thickened and flared; 14–37 rounded axial ribs on penultimate whorl; ventrolateral varix a moderate rib at 200–270 °; 5 spiral cords on spire, 5– 9 above periphery on last whorl; often a broad brown band from periphery to shoulder. E India, SE Asia to Borneo and Java. COI GenBank HE 680209 –680219, AM 932765. Material examined. 106 lots. Shell (Fig. 8): H = 31.3–57.2 mm. Shape relatively broad, elongated conical (H/B = 1.78–1.91, SH = 2.20– 2.59); decollate, 5–7 whorls remaining; spire whorls rounded, suture distinct; spire profile straight to slightly convex; periphery rounded, not marked by cord or keel; solid. Adult lip strongly thickened and flared; apertural margin planar in side view; strong anterior projection adjacent to canal. Sculpture on spire of slightly opisthocline curved (opisthocyrt) axial ribs, occasionally bifurcating posteriorly (adapically), usually weaker and more distant on final 0.3 whorl; ribs prominent, rounded, interspaces 1–2 times width of ribs, 14–37 ribs on penultimate whorl; occasionally 1–2 varices on spire; spire whorls with 5 spiral cords, intersecting with axial ribs to give reticulate sculpture, increasing by interpolation of narrower cords to 6–9 spiral cords above periphery on final whorl; base with 10–13 cords, only slightly narrower than those above periphery and not separated from them. Ventrolateral varix a slightly to moderately thickened rib at 200–270 °. Surface with fine spiral microstriae on periostracum. Colour: fawn, usually a broad dark brown band from shoulder to periphery and another on base; aperture white, pale brownish within. Animal (Fig. 2 B): Head and base of tentacles grey with pale yellow spots; snout dark grey with 3 transverse bands of orange red, the 2 posterior bands wavy or interrupted and with a broad black band between; eyes with black surround and red ring; tentacles pinkish grey, darker at base; sides of foot dark grey with red spots and red margin; mantle cream (red colour fading to dark pink in ethanol-preserved specimens.) Range (Fig. 9): Bay of Bengal, Malay Peninsula, S Vietnam, Sumatra, Java, Borneo. Records: India: Pichavaram, Tamilnadu (Sreenivasan & Natarajan 1986; Subba Rao 2003); Godavari Estuary, Andhra Pradesh (Subba Rao, 2003); Port Canning, West Bengal (NHMUK 1893.12.2.119). Bangladesh: R. Ganges (ZMB). Burma: Moulmein, Salween R. (NHMUK 1954.6.2.2). Thailand: Ko Nok, Satun Prov. (NHMUK; MNHN; RMNH; AM C.079263; USNM 777316); Kanchanadit, Surat Thani Prov. (NHMUK 20100436; USNM 828815); Ban Tamrua, Chonburi Prov. (USNM 776723). Malaysia: Klang, Selangor (NHMUK; USNM 661023; ANSP 267498); Kuantan, Pahang (AM C. 124490); Simunjan, Sarawak (ZMB 112.764). Singapore: Pulau Ubin (NHMUK 20130239). Cambodia: Kampot (Morlet 1889). Vietnam: Vung Tau (ANSP 330753); Long Hai (NHMUK); Khanh Hoa (Thach 2005). Indonesia: Pulau Weh, Sumatra (RMNH); Pulau Bangka (AM C.009036); Tanjung Priok, Java (RMNH); Pasuruan, Java (RMNH); Banjarmasin, Kalimantan (RMNH); 14 km N Mahakan R., Kalimantan (AM C. 410364). Lamarck (1822) gave the type locality as “Mers de Timor” but, at this early date, this might refer to a wider region than the island of Timor. There are in addition four museum records from Timor (RMNH; ANSP 18055; ZMB), but all are old and are considered unreliable; the locality may have been copied from Lamarck, or ships returning to Europe from Timor may also have acquired specimens from Java or elsewhere. Three species of Potamididae were listed from the mangroves of Timor by Studer (1889), but C. obtusa was not among them. The Australian C. reidi and C. anticipata have sometimes been misidentified as C. obtusa (see their respective Synonymies). Houbrick (1986: 284), while distinguishing all three, also noted that C. obtusa “does not appear to be common in Australia, although there are some reliable records from Queensland”. In fact there is no evidence that C. obtusa occurs in Australia. Among other authors, von Martens (1897 a, b) included Australia and Madagascar in the recorded distribution; Brandt (1974) included Madagascar, Philippines and Australia; and Poutiers (1998) also mentioned Madagascar. Its incorrect occurrence in Madagascar can be traced to the record by Kiener (1841 –1842), who was later quoted by von Martens (1880) and Dautzenberg (1923, 1929). The distribution in India has been said to include the Andaman and Nicobar islands (Subba Rao 2003; Subba Rao et al. 1992), but this is unsubstantiated; occurrence of Cerithidea there could refer to C. andamanensis. Records of shells outside its natural range could possibly be a consequence of its widespread use as food in Southeast Asia. Habitat and ecology. In Rhizophora and Bruguiera forest, Avicennia parkland and landward fringes of mangrove forests, on firm mud and on trunks and stilt roots up to 1–1.5 m above the ground; in fully marine and estuarine areas. It has been recorded from the Rajang River at Sibu, Sarawak (Hamli et al. 2013), 60 km from the open sea. Several authors have observed that C. obtusa rests on branches, roots and trunks of trees, and descends to feed on firm mud below as the tide recedes, before climbing again during the ebb tide (Sreenivasan & Natarajan 1986; Ahmed & Mahmud 2007); according to Berry (1972) descent to the forest floor occurs during neap tides in Malaysia. In Malaysia and Thailand the animals are often found close to the ground on the bases of trees and on firm mud among tree roots (pers. obs.), but presumably climb higher to avoid submersion during spring tides. In Bangladesh the snails are said to be often found partly buried in the mud with the top of the spire projecting (Ahmed & Mahmud 2007). Densities of 2–10 /m 2 have been recorded in Sundarbans (Subba Rao et al. 1983, 1995; Dey 2006). It was rare in a mangrove forest at Phuket, Thailand, perhaps as a result of collection for food by local people (Frith, Tantanasiriwong & Bhatia 1976). According to the growth rate estimated by Sreenivasan & Natarajan (1986) in India, 13 mm is achieved at the end of the first year and maximum size of 26 mm after 4 years. Houlihan (1979) measured an air/water respiration rate ratio of 5.5 at 28 °C and reported 50 % death after submersion for 48 h, also at 28 °C. Remarks. The sculpture of the shell is quite variable (as observed by Sowerby 1866: fig. 4 a, b), specimens from India and Bangladesh (Fig. 8 G, H) usually having finer sculpture than the typical form from the western Pacific (Fig. 8 I), but both can be found, together with intermediates, on the west coast of the Malay Peninsula (Fig. B–F, J–M). A phylogeographic break between Indian and Pacific oceans is common in shallow-water marine organisms (e.g. Reid et al. 2006; Carpenter et al. 2011) and in some cases sister species occur in each ocean (Gaither & Rocha 2013), but the existence of intermediates suggests conspecificity in the case of C. obtusa. In addition, the distinctive coloration of the body is the same in both forms; Subba Rao (2003: 136) reported “alternating broad black and red bands” on the head in India. Samples often show size dimorphism of small and large shells (e.g. Fig. 8 G, H), which presumably represent male and female individuals. The species is commonly used for food in Southeast Asia and Indonesia (Brandt 1974; Poutiers 1998; Hamli et al. 2013). Cerithidea obtusa is a member of a large clade of Cerithidea species (excluding the C. balteata group and C. weyersi), but within this its relationships are not clearly resolved in molecular phylogenies (Reid et al. 2013; Fig. 1). The shell is similar to that of C. anticipata, but is larger, broader and more robust. It is also close to C. reidi, which is of similarly large size, but paler in colour. 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Cerithidea moerchii (A. Adams in G.B. Sowerby II, 1855) (Figures 2 E, 14 B, 15 J–AA) Cerithium mörchii A. Adams in G.B. Sowerby II, 1855: 888, pl. 186, fig. 283 (no locality, here corrected to Yaeyama Is, Okinawa Pref., Japan; 2 syntypes NHMUK 20060321, Fig. 15 U, seen). Cerithidea moerchii — Sowerby, 1866: sp. 18, pl. 3, fig. 18. Kuroda, 1928: 38 (as mörchii). Hirase, 1934: 53, pl. 84, fig. 8 (as mörchii). Kuroda & Habe, 1952: 44 (as morchii). Higo et al., 1999: 87 (as morchii). Reid et al., 2013: figs 1 (phylogeny), 2 (map). Potamides (Cerithidea) moerchii — Tryon, 1887: 163, pl. 33, fig. 68 (as mörchi). Iwakawa, 1919: 76 (as morchi). Cerithium (Cerithidea) moerchii — Kobelt, 1893: 154, pl. 29, fig. 3 (Cecalupo 2005: pl. 32, fig. 12). Cerithidea (Cerithidea) moerchii — Oyama, 1959: figs 4, 11 (as mörchii). Cerithidea rhizophorarum moerchii — Habe & Kosuge, 1967: 34, pl. 13, fig. 16 (as morchii). Habe, 1968: 38, pl. 11, fig. 19 (as rhizoghorarum morchi). Higo, 1973: 59 (as morchii). Okutani, 1986: 80, fig. (as morchi). Higo & Goto, 1993: 105 (as morchii). Cerithidea obtusa var. β rhizophorarum — Crosse & Fischer, 1864: 332 (not A. Adams, 1855). Cerithidea rhizophorarum — Dautzenberg & Fischer, 1905: 134 –135 (in part, includes C. rhizophorarum; as rhizoporarum). Hirase, 1934: 53, pl. 84, fig. 7 (as rhizoporarum). Kuroda & Habe, 1952: 44. Habe & Kosuge, 1967: 34, pl. 13, figs 13–15. Habe, 1968: 38, pl. 11, fig. 23. Kuroda et al., 1971: 101, pl. 16, figs 1, 2. Higo, 1973: 59. Wells, 1985: 145, pl. 1 H. Okutani, 1986: 80, figs. Fan 1989: 38, pl. 5, fig. 25. Choe, 1992: 309 –310, pl. 64, fig. 101 a, b. Higo & Goto, 1993: 105. Higo et al., 1999: 87. Lee & Min, 2002: 103. Ma, 2004: 37, pl. 016C (not 016A). Min et al., 2004: 129, fig. 217-1, 217 - 2. Kojima et al., 2006: 525 –535, fig. 2 c, d. Reid et al., 2008: 680 –699, figs 1, 2 (phylogeny). Zhang, 2008: 59, text fig. Li & Huang, 2012: 152, text fig. (All not A. Adams, 1855.) Potamides (Cerithidea) rhizophorarum — Iwakawa, 1919: 76 (not A. Adams, 1855). Cerithidea (Cerithidea) rhizophorarum — Oyama, 1959: figs 5, 12. Fukuda, 1996: 21 –22. Fukuda, 2000: 106 –110, 135, fig. 5.4 a (shell), fig. 5.5 a (living animal). Hasegawa, 2000: 133, pl. 66, fig. 1 a, b. Cecalupo, 2006: 136 (in part, includes C. rhizophorarum, C. decollata, C. weyersi). Thach, 2007: 60, pl. 9, fig. 183. Hong et al., 2010: 258, fig. 4 c. Kimura & Fukuda, 2012: 29, text figs. a, b, d, e (shells), c (living animal). (All not A. Adams, 1855.) Cerithidea rhizophorarum var. freytagi — Dautzenberg & Fischer, 1905: 135 (in part, includes C. rhizophorarum; as freitagi; not Kobelt, 1893). Cerithidea obtusa — Grabau & King, 1928: 223 –224, pl. 9, fig. 95 (not Lamarck, 1822). Cerithidea (Cerithideopsis) scalariformis — Cecalupo, 2005: 318, pl. 32, fig. 12 (not Say, 1825). Cecalupo, 2006: 104 (in part, includes C. balteata; not Say, 1825). Cerithidea ornata — Li & Huang, 2012: 152, fig. (as ornate; not Sowerby, 1855). Taxonomic history. For more than a century this species has been most familiar under the incorrect name C. rhizophorarum (here shown to be a distinct species, endemic to the Philippines). The name C. moerchii was not, however, forgotten. In the Ryukyu Islands and Taiwan the shells are small with coarse ribs, and C. moerchii was based on this form. Early Japanese authors identified the larger mainland form as C. rhizophorarum and used C. moerchii for the smaller island form, considering it to be a distinct species (Iwakawa 1919; Kuroda 1928; Hirase 1934; Kuroda & Habe 1952; Oyama 1959; Higo et al. 1999) or subspecies (Habe & Kosuge 1967; Habe 1968; Higo 1973; Okutani 1986; Higo & Goto 1993). More recently, C. moerchii has been considered only a “geographical form” (Hasegawa 2000: 133) and this was supported by the lack of genetic differentiation between samples from mainland Japan and the Ryukyu Islands (Kojima et al. 2006; Reid et al. 2008, 2013). The name C. rhizophorarum has since been widely applied in Japan, China, Korea and Vietnam (e.g. Ma 2004; Min et al. 2004; Thach 2007; Zhang 2008; Kimura & Fukuda 2012; Li & Huang 2012), but C. moerchii has now been reinstated as the valid name for this species (Reid et al. 2013). The name moerchii first appeared in Sowerby’s (1855) monograph of Cerithium, with its authorship attributed to A. Adams. In his introduction, Sowerby (1855: 848) acknowledges Adams “for his kindness in permitting me the full use of his manuscripts on the present genus, of which he was preparing a monograph”. As noted earlier (see Taxonomic history of C. balteata), Sowerby employed only the single genus Cerithium for all members of the modern families Cerithiidae, Potamididae, Batillariidae and others, whereas Adams employed a finer division, including the genus Cerithidea. Adams’ (1855) monograph of Cerithidea was in fact published just before Sowerby’s work appeared (Petit 2009). It is unclear whether Sowerby saw this completed text or only its manuscript, but he certainly relied on Adams’ work, because he used most of Adams’ new names and copied some of his descriptions. It is curious, therefore, that Cerithium moerchii, attributed to Adams, appears only in Sowerby’s monograph and not in that of Adams. In one other case (see discussion of Cerithium ornatum in Taxonomic history of C. balteata) Sowerby apparently replaced one of Adams’ names to avoid homonymy in the genus Cerithium, but this does not apply to Cerithium moerchii. The text does not disclose why this name was chosen, honouring Mörch. The two syntypes are from the Cuming Collection (NHMUK) and Cuming and Mörch are known to have exchanged specimens from the Philippines (Dance 1986: 158), which may provide an explanation. Authorship of this species is here attributed to ‘Adams in Sowerby’ (as also by Petit 2009). The original description gave no locality, but simply noted that it was “a northern species” (Sowerby 1855: 888), yet in a subsequent monograph the locality of the same sample was given as Mindanao, Philippines (Sowerby 1866). No original label survives with the type material and the species is not known to occur in the Philippines. Shells exactly similar to the distinctive syntypes have been seen only from Ishigaki and Iriomote Islands in the Yaeyama group, so the type locality is here corrected accordingly. The change from mörchii to moerchii is a mandatory correction of an incorrect original spelling (ICZN 1999: Art. 32.5.2.1). Diagnosis. Shell: periphery rounded, spire whorls well rounded; spire profile straight; aperture slightly thickened and flared, sinuous apertural margin; 14–34 rounded axial ribs on penultimate whorl, ribs usually weak or absent after ventrolateral varix; ventrolateral varix a wide rib at 170–230 (150–330)°; 0–6 varices on spire; 5–10 spiral cords above periphery, often producing cancellate effect; banded or lined pattern. Southern Japan, South Korea, China, Vietnam. COI GenBank HE 680239 – HE 680293, AM 932773 – AM 932775. Material examined. 68 lots. Shell (Fig. 15 J–AA): H = 16.1–46.6 mm. Shape elongated conical (H/B = 2.08–2.94; SH = 3.0– 4.04); decollate, 5–8 (13) whorls remaining; spire whorls moderately to well rounded, suture distinct; spire profile straight; 0–6 thick pale varices on spire; periphery rounded; moderate thickness. Adult lip slightly thickened and flared; apertural margin sinuous in side view; weak anterior projection adjacent to canal. Sculpture on spire of straight to slightly opisthocline axial ribs, becoming slightly curved (opisthocyrt) on last whorl, ribs rounded, interspaces 1–1.5 times width of ribs, 14–23 (34) ribs on penultimate whorl; axial ribs usually weak or absent after ventrolateral varix, but rarely up to 8 are present; spiral sculpture of 5 primary cords on spire whorls, of which anterior cords usually divide on later whorls to give 6–9 cords on penultimate whorl and up to 10 above periphery on final whorl, posterior cord adjacent to suture strongest and bearing small nodules (corresponding with, or up to twice as numerous, as axial ribs); intersection of axial ribs and spiral cords often giving cancellate effect; base with 8–10 ridges, merging with cords above rounded periphery. Ventrolateral varix a wide rib at 170–230 (150–330)°; 0– 6 wide varices on spire. Surface with fine spiral microstriae on periostracum; surface dull when worn. Colour: white, fawn or pale brown, usually with dark brown spiral lines and bands; often a dark band at shoulder and on base, or single broad band from shoulder to periphery, or only spiral grooves dark brown, varices white; bands show through in aperture. Animal (Fig. 2 E): Head pale brown with cream spots; snout brown to black with cream spots, darker towards tip; tentacles fawn with dark rings, black at base, pinkish around eye; sides of foot grey to blackish brown with cream spots; sole of foot grey; mantle cream (ethanol-preserved specimens similar). Range (Fig. 14 B): S Japan to Vietnam. Records: Japan: Matsushima Bay, Miyagi Pref. (ANSP 241064); Yokohama (ANSP 49910); Yoshida R., Oiehama, Kasaoka city, Hiroshima Pref. (NHMUK 20130265); Imaizumi, Kagoshima Pref. (USNM 363689); Jigyo, Chuo-ku, Fukuoka Pref. (ANSP 418823); Ura, Tatsugou-cho, Amami city, Kagoshima Pref. (NHMUK 20130271); Waku-gawa, Gabui, Nakijin-son, Okinawa (NHMUK); Funaura Bay, Iriomote I. (NHMUK 20130270). S Korea: Suncheon Bay (Hong, Choi & Tsutsumi 2010); Jeju I. (Noseworthy et al. 2007). China: “throughout Chinese coasts” (Ma 2004: 37); Tolo Harbour, Hong Kong (NHMUK 20130263); Hainan (ZMSP). Taiwan: Tamshui R., Bali (NHMUK 20070387); Kenting National Park (Lee & Chao 2004). Vietnam: Tuan Chau I., Ha Long Bay (NHMUK); Lo R., Nha Trang (NHMUK); Long Hai (Thach 2005). An old record from Hakodate, in the south of Hokkaido (USNM unregistered) is unconfirmed. The localities Borneo and Labuan [Sabah] on specimens in NHMUK are considered to be incorrect. Habitat and ecology. On firm or sandy mud and stones of tidal flats, in the open or under mangroves; often at river mouths and in tidal creeks; among salt-tolerant grasses and Phragmites reeds behind mud flats; sometimes on trunks of mangroves including Kandelia and Rhizophora. A number of authors have described the habitat and habits of this species. In Hong Kong it has been reported as common throughout mangrove stands from seaward to landward, both on the substrate and up to 1.0 m on trunks, where it attaches by mucus when dry (McMahon 1985); at upper tidal levels it also occurs on grasses, sand and rocks in the absence of mangroves (Wells 1985). In Korea, Hong et al. (2010) found it on mud flats among Suaeda bushes. Most observations have been made in southern Japan. The northernmost mangrove forest in the western Pacific is situated at Atago River in southern Kyushu, where the snails are distributed both on trunks of Kandelia and on the tidal flats of sandy mud. The animals feed on the substrate during low tide (Maki, Ohtaki & Tomiyama 2002), but climb trees to avoid submersion at high tide, especially from August to January or in the winter (Ohtaki et al. 2002; Takeuchi, Ohtaki & Tomiyama 2007). The snails not only climb mangroves trees, but also on reeds and rocks (Matsumura 2009). The smallest snails occur only on unvegetated mud flats seaward of salt marsh and mangroves, while adults are more widely distributed (Ohtaki, Maki & Tomiyama 2001; Wada & Nishikawa 2005; Takeuchi et al. 2007). In a population at Matsushima Bay, beyond the northern limit of mangroves, isotopic analyses showed that the major food source was detritus from macroalgae and seagrass, not benthic microalgae (Doi et al. 2009). In Japan, pseudocopulation takes place from June to August during daytime and nighttime low tides and occurs by males mounting the shells of females (Ohtaki et al. 2001; Takeuchi et al. 2007; Ota, Kawai & Hashimoto 2013). During the process a 3 -mm long, spindle-shaped spermatophore is transferred (Onoda et al. 2010). Spawning has been observed from June to August (Wada & Nishikawa 2005; Ota et al. 2013) and was described in detail by Habe (1955). A mass of gelatinous egg strings, about 20 mm wide and 15 mm deep, is deposited in an excavation in the mud. Individual eggs are 0.35 mm in diameter. Houbrick (1984: 12) interpreted this relatively large egg size as “suggesting direct development”, but Kimura et al. (2002) mentioned the existence of planktonic larvae. Kojima et al. (2006) claimed a ‘planktotrophic stage’ of 12–20 days (quoting Kimura et al. 2002 and unpublished work). Ota et al. (2013) also observed that swimming larvae hatched from the eggs. (In a more detailed account of the western Atlantic potamidid Cerithideopsis pliculosa, Miura, Frankel & Torchin 2011 described a shell diameter of 0.3 mm at hatching and observed swimming-crawling, planktotrophic larvae that metamorphosed after 6–13 days, lending indirect support to this claim.) Ota et al. (2013) described the population dynamics of C. moerchii on a bare tidal flat in E Shikoku, where density varied from 14–79 /m 2 over 3 years. Recruitment occurred in late summer and early autumn, and thickened apertural lips (implying reproductive maturity) developed after the third year. Older cohorts occupied lower positions (around mean tide level) in summer; recruitment occurred at these lower levels and was followed by a gradual upward shift. Remarks. This species has a wide distribution from the temperate zones of central Japan, Korea and northern China to the tropical areas of Vietnam (Fig. 14 B). Correspondingly, it has a wide range of size and shell morphology. Several distinctive regional forms can be recognized, but there are intergradations between them: (1) Southern China, Vietnam (Fig. 15 J–N): less rounded whorls; numerous narrow axial ribs (usually 19–27); white varices often present on spire. (2) Mainland Japan (Fig. 15 O–T): large shell; rounded whorls; strong relatively broad axial ribs (usually 19–23); spire varices usually absent; dark or banded pattern. (3) Amami Islands, Okinawa, Yaeyama Islands, Taiwan (Fig. 15 U–AA): small shell; rounded whorls; strong relatively broad axial ribs (14–18); spire varices usually absent; most primary cords become divided, giving 7– 10 fine spiral ridges on last whorl; axial ribs continue after ventrolateral varix; unbanded but often with dark spiral grooves. In twentieth century literature, forms (2) and (3) have usually been distinguished as separate species (or subspecies), respectively C. ‘rhizophorarum’ and C. moerchii (see Taxonomic history, above). Despite the distinctive appearance of shells from the southern islands, molecular investigation has failed to find any significant differentiation in COI sequences between these and samples from mainland Japan (Kojima et al. 2006, Reid et al. 2008, 2013). Likewise, Kamimura et al. (2010) found no significant differentiation of this gene among four populations from mainland Japan and Korea. Examination of large series of shells from throughout the range also shows that there is intergradation among the forms (Fig. 15 J–AA). The larger forms of C. moerchii from China, Korea and Japan are very similar in appearance to C. rhizophorarum, which is restricted to the Philippines, explaining the longstanding confusion of these taxa. Cerithidea moerchii has more rounded whorls, a straight spire profile and shows a wider range of shell patterns, including forms with narrow spiral bands. Forms from China and mainland Japan are more distinctly cancellate than C. rhizophorarum, especially when axial ribs are narrow, and the broad axial ribs of shells from the Ryukyus give a rugose appearance unlike any specimens of C. rhizophorarum. The latter species has flattened whorls, a pupoidal outline, the axial ribs are always narrow, and narrowly banded colour forms have not been seen (Fig. 15 A–I). It is predicted that these two will prove to be sister species when their status is tested with molecular data from C. rhizophorarum. In the COI phylogenetic tree (Reid et al. 2013; Fig. 1) C. rhizophorarum was not represented, so C. moerchii appears as sister to C. tonkiniana. Occasional specimens show an unusually elongate shell with slightly irregular whorls (Fig. 15 S); such shells appear to be the result of abnormal growth, possibly associated with parasitism by trematodes or other environmental influence. Similar malformations have been seen in Cerithideopsilla species from the northwestern Pacific region. Owing to habitat loss, this species is listed as ‘vulnerable’ in Japan by Fukuda (1996) and as ‘near threatened’ by Kimura & Fukuda (2012)., Published as part of Reid, David G., 2014, The genus Cerithidea Swainson, 1840 (Gastropoda: Potamididae) in the Indo-West Pacific region, pp. 1-65 in Zootaxa 3775 (1) on pages 52-55, DOI: 10.11646/zootaxa.3775.1.1, http://zenodo.org/record/285731, {"references":["Sowerby, G. B. II (1855) Monograph of the genus Cerithium, Adanson. In: Sowerby, G. B. II. (Ed.), Thesaurus Conchyliorum, or Figures and Descriptions of Recent Shells. Vol. 2. part 16. Sowerby, London, pp. 847 - 899.","Sowerby, G. B. II (1866) Monograph of the genus Cerithidea. In: Reeve, L. A. (Ed.), Conchologia Iconica. Vol. 15. L. A. Reeve, London, pls 1 - 4.","Kuroda, T. (1928) Catalogue of the Shell-bearing Mollusca of Amami-Oshima (Oshima, Osumi). Kagoshima-Ken Educational Investigation Committee, Kagoshima, 126 pp. [in Japanese]","Hirase, S. 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Eru Malacological Publisher, Osaka, 3 + 22 + 693 + 13 + 149 pp.","Crosse, H. & Fischer, P. (1864) Faune malacologique de Cochinchine. - Premier supplement. Journal de Conchyliologie, 12, 322 - 338.","Adams, A. (1855) A monograph of Cerithidea, a genus of Mollusca, with descriptions of several new species, from the collection of High Cuming, Esq,: to which are added, descriptions of two new species of Colina, and one of Donax. Proceedings of the Zoological Society of London, 1854, 83 - 87. http: // dx. doi. org / 10.1111 / j. 1469 - 7998.1854. tb 07236. x","Dautzenberg, P. & Fischer, H. (1905) Liste des mollusques recoltes par M. le Capitaine de Fregate Blaise au Tonkin, et description d'especes nouvelles. Journal de Conchyliologie, 53, 85 - 234.","Kuroda, T., Habe, T. & Oyama, K. (1971) The Sea Shells of Sagami Bay. Maruzen, Tokyo, xix + 741 + 489 + 51 pp.","Wells, F. E. (1985) The Potamididae (Mollusca: Gastropoda) of Hong Kong, with an examination of habitat segregation in a small mangrove system. In: Morton, B. & Dudgeon, D. (Eds) Proceedings of the Second International Workshop on the Malacofauna of Hong Kong and Southern China, Hong Kong. Hong Kong University Press, Hong Kong, pp. 139 - 154.","Fan, C. (Ed.) (1989) Mollusca of Huanghai and Bohai. Nongye Chubanshe, Agricultural Publishing House, Beijing, 309 pp.","Choe, B. L. (1992) Mollusca (II). Illustrated Encyclopaedia of Fauna and Flora of Korea. Vol. 33. Ministry of Education, Republic of Korea, 860 pp.","Lee, J. - S. & Min, D. - K. (2002) A catalogue of molluscan fauna in Korea. Korean Journal of Malacology, 18, 93 - 217.","Ma, X. (2004) Order Mesogastropoda. In: Qi, Z. (Ed.), Seashells of China. China Ocean Press, Beijing, pp. 31 - 81.","Min, D. - K., Lee, J. - S., Koh, D. - B. & Je, J. - G. (2004) Mollusks in Korea. Min Molluscan Research Institute, Seoul, 566 pp.","Kojima, S., Kamimura, S., Iijima, A., Kimura, T., Kurozumi, T. & Furota, T. (2006) Molecular phylogeny and population structure of tideland snails in the genus Cerithidea around Japan. Marine Biology, 149, 525 - 535. http: // dx. doi. org / 10.1007 / s 00227 - 005 - 0183 - 2","Reid, D. G., Dyal, P., Lozouet, P., Glaubrecht, M. & Williams, S. T. (2008) Mudwhelks and mangroves: the evolutionary history of an ecological association (Gastropoda: Potamididae). Molecular Phylogenetics and Evolution, 47, 680 - 699. http: // dx. doi. org / 10.1016 / j. ympev. 2008.01.003","Zhang, S. (2008) Atlas of Marine Mollusks in China. Hai yang chu ban she, Beijing, iv + 383 pp.","Li, R. & Huang, Z. (2012) Class Gastropoda. In: Huang, Z. & Liu, M. (Eds.), The Living Species and their Illustrations in China's Seas (Part II). An Illustrated Guide to Species in China's Sea. Vol. 4. Ocean Press, Beijing, pp. 102 - 279.","Fukuda, H. (1996) Gastropoda. In: Hanawa, S. & Sakuma, H. (Eds.), Present Status of Estuarine Locales and Benthic Invertebrates Occurring in Estuarine Environment in Japan. WWF Japan Science Report, 3, 11 - 63.","Fukuda, H. (2000) Gastropods I: general view. In: Sato, M. (Ed.), Life in Ariake Sea: Biodiversity in Tidal Flats and Estuaries. Kaiyu-sha, Tokyo, pp. 213 - 252.","Hasegawa, K. (2000) Family Potamididae. In: Okutani, T., (Ed.), Marine Mollusks in Japan. Tokai University Press, Tokyo, pp. 133 - 134.","Cecalupo, A. (2006) Elenco della famiglia Cerithiidae Ferussac, 1822 (Prosobranchia). Nomenclatura delle specie conosciute o poco note e relativa revisione sistematica. Attuali. Vol. II. Quaderni della Civica Stazione Idrobiologica di Milano, 27, 1 - 373.","Thach, N. N. (2007) Recently Collected Shells of Vietnam. L'Informatore Piceno, Ancona, Italy, 384 pp.","Hong, J. - S., Choi, J. - W. & Tsutsumi, H. (2010) Concluding remarks on the joint survey of macrobenthic fauna on Suncheon Tidal Flats by the participants of \" Korea and Japan Joint Symposium on Biology of Tidal Flats 2009 \". Plankton & Benthos Research, 5 (suppl.), 255 - 263. http: // dx. doi. org / 10.3800 / pbr. 5.255","Grabau, A. W. & King, S. G. (1928) Peking Society of Natural History, Handbook no. 2. Shells of Peitaiho. Edition 2. Peking, Peking Laboratory of Natural History, vi + 279 pp.","Lamarck, J. B. P. A. de (1822) Histoire Naturelle de Animaux sans Vertebres. Vols 6 (2), 7. Paris, 232 pp., 711 pp.","Say, T. (1825) Descriptions of some new species of fresh water and land snails of the United States. Journal of the Academy of Natural Sciences of Philadelphia, 5, 119 - 131.","Petit, R. E. (2009) George Brettingham Sowerby, I, II & III: their conchological publications and molluscan taxa. Zootaxa, 2189, 1 - 218.","Dance, S. P. (1986) A History of Shell Collecting. E. J. Brill, Leiden, vi + 265 pp.","Noseworthy, R. G., Lim, N. - R. & Choi, K. - S. (2007) A catalogue of the mollusks of Jeju Island, South Korea. Korean Journal of Malacology, 23, 65 - 104.","Lee, S. - C. & Chao, S. - M. (2004) Shallow-water marine shells from Kenting National Park, Taiwan. Collections and Research, 17, 33 - 57.","Thach, N. N. (2005) Shells of Vietnam. Conchbooks, Hackenheim, 338 pp.","McMahon, R. F. (1985) Interspecific relationships between morphometric parameters and the vertical distribution patterns of seven species of turbinate gastropods on mangrove trees in Hong Kong. In: Morton, B. & Dudgeon, D. (Eds.), Proceedings of the Second International Workshop on the Malacofauna of Hong Kong and Southern China, Hong Kong. Hong Kong University Press, Hong Kong, pp. 199 - 215.","Takeuchi, M., Ohtaki, H. & Tomiyama, K. (2007) Reproductive behavior of the dioecious tidal snail Cerithidea rhizophorarum (Gastropoda: Potamididae). American Malacological Bulletin, 23, 81 - 87. http: // dx. doi. org / 10.4003 / 0740 - 2783 - 23.1.81","Matsumura, I. (2009) Interesting behaviour of Cerithidea rhizophorarum (Gastropoda: Potamididae) attaching to reed grass. Chiribotan, 39, 153 - 155. [in Japanese]","Ohtaki, H., Maki, E. & Tomiyama, K. (2001) Seasonal changes in the distribution and mating behavior of Cerithidea rhizophorarum (Gastropoda: Potamididae). Venus, 60, 199 - 210.","Wada, K. & Nishikawa, C. (2005) Habitat use by the potamidid gastropod Cerithidea rhizophorarum living in an estuarine salt marsh. Japanese Journal of Benthology, 60, 23 - 29. [in Japanese]","Doi, H., Matsumasa, M., Fujikawa, M., Kanou, K. Suzuki, T. & Kikuchi, E. (2009) Macroalgae and seagrass contribution to gastropods in sub-tropical and temperate tidal flats. Journal of the Marine Biological Association of the United Kingdom, 89, 399 - 404. http: // dx. doi. org / 10.1017 / s 0025315408002683","Ota, N., Kawai, T. & Hashimoto, A. (2013) Recruitment, growth, and vertical distribution of the endangered mud snail Cerithidea rhizophorarum A. Adams, 1855: implications for its conservation. Molluscan Research, 33, 87 - 97. http: // dx. doi. org / 10.1080 / 13235818.2013.782789","Onoda, G., Suzuka, T., Takeuchi, Y., Konagai, T. & Tomiyama, K. (2010) Spermatopohore transfer in the dioecious tidal snail Cerithidea rhizophorarum (Gastropoda: Potamididae). Venus, 68, 3 - 4.","Habe, T. (1955) Spawning of Cerithidea djadjariensis and C. rhizophorarum. Venus, 18, 204 - 205.","Houbrick, R. S. (1984) Revision of higher taxa in genus Cerithidea (Mesogastropoda: Potamididae) based on comparative morphology and biological data. American Malacological Bulletin, 2, 1 - 20.","Kimura, T., Fujioka, E., Kimura, S. & Aoki, S. (2002) A comparative morphology of eggs and larvae of nine gastropods in tidal flat and marsh of the reed. Venus, 61, 114 - 115. [in Japanese]","Miura, O., Frankel, V. & Torchin, M. E. (2011) Different developmental strategies in geminate snails, Cerithideopsis californica and C. pliculosa, across the Isthmus of Panama. Journal of Molluscan Studies, 77, 255 - 258. http: // dx. doi. org / 10.1093 / mollus / eyr 012","Kamimura, S., Itoh, H., Ozeki, S. & Kojima, S. (2010) Molecular diversity of Cerithidea gastropods inhabiting Suncheon Bay, and the Japanese and Ryukyu Islands. Plankton & Benthos Research, 5 (suppl.), 250 - 254. http: // dx. doi. org / 10.3800 / pbr. 5.250"]}

Cerithidea reidi Houbrick, 1986 (Figures 2 G, 10 Q–V, 11) Cerithidea obtusa — Allan, 1950: 87, pl. 17, fig. 19. Hinton, 1978: 7, fig. Wells, 1981: 254. Wells & Bryce, 1986: 54, pl. 11, fig. 128. (All not Lamarck, 1822.) Cerithidea reidi Houbrick, 1986: 280 –286, figs 1 –4, 9 and 11–13 (radula), 16 (map) (Willies Creek, N of Broome, Western Australia; holotype WAM 3380 - 84; 1 paratype AM C. 144144; 10 paratypes USNM 828823 seen). Wells & Bryce, 1988: 54, pl. 11, fig. 128. Cecalupo, 2006: 135, 233– 234, pl. 57, fig. 14 (in part, includes C. anticipata, C. dohrni). Willan, 2013: 75 –77, fig. 7. Reid et al., 2013: figs 1 (phylogeny), 2 (map). Cerithidea (Cerithidea) reidi — Wilson, 1993: 133, pl. 15, fig. 17 a, b. Taxonomic history. Until clarified by Houbrick (1986), there was confusion about the identity of the two Australian Cerithidea species. Until then, this endemic Western Australian species had been figured several times under the name ‘ C. obtusa ’. This species was named for the author by Richard Houbrick; he inscribed a reprint with the epigram “to David – immortalized until synonymized”! Diagnosis. Shell: periphery rounded or weakly angled; aperture flared, anterior canal and apertural projection well developed; 19–22 axial ribs on penultimate whorl; ventrolateral varix a slightly enlarged rib at 190–250 °; 5 broad spiral cords on spire, 5–10 above periphery on last whorl; whitish fawn with orange brown spiral grooves and lilac aperture. Northern Western Australia. COI GenBank HE 680236 –680238. Material examined. 15 lots. Shell (Fig. 10 Q–V): H = 46.0–60.0 mm (to 61 mm, Houbrick 1986). Shape elongated conical (H/B = 1.95– 2.38, SH = 2.36–2.80); decollate, 6–8 whorls remaining; spire whorls rounded, suture distinct; spire profile straight to slightly convex; periphery rounded or weakly angled; moderately solid. Adult lip flared, moderately thickened; apertural margin planar in side view; strong anterior projection adjacent to deep notch of anterior canal. Sculpture on spire of straight to slightly curved (opisthocyrt) axial ribs, often bifurcating or trifurcating posteriorly (adapically) to give a row of granules or short axial wrinkles at suture; ribs prominent, rounded, ribs and interspaces of equal width, 19–22 ribs on penultimate whorl; becoming weak on last whorl (especially before ventrolateral varix); continuing as closely spaced, sometimes rugose, axial wrinkles after ventolateral varix, fine on base; spire whorls with 5 strap-like spiral cords, often forming weak nodules where they cross axial ribs, usually increasing by interpolation of narrower threads to give 5–10 spiral cords and threads above periphery on final whorl; base with 9–14 narrow ridges, peripheral ridge narrower than primary cords above, but is larger than other basal ridges. Ventrolateral varix a slightly enlarged rib at 190–250 °. Surface with faint spiral microstriae on periostracum. Colour: whitish fawn, palest at suture, spiral grooves orange brown to lilac grey; aperture mauve, lilac or pale pinkish brown, spiral lines showing through faintly, lip white to cream. Animal (Fig. 2 G): Head and base of tentacles pinkish fawn with cream spots; anterior half of snout blackish grey with scattered yellow spots; tentacles cream, black annular rings; sides of foot fawn with darker specking and paler patches, small yellow spots at margin; sole of foot pale grey; mantle pinkish grey. Range (Fig. 11): N Western Australia (WA). Records: Australia: Gales Bay, Exmouth Gulf, WA (Houbrick 1986: WAM 2434 - 84); Barrow I., WA (USNM 694228); Willies Creek, N of Broome, WA (NHMUK 20130246; USNM 828823); Buccaneer Arch., WA (AM C. 042225); Port Warrender, Admiralty Gulf, WA (WAM S 46091; Houbrick 1986). Habitat and ecology. Near Broome the species is found on trunks of Ceriops bushes and trees, close to deep tidal creeks crossing bare salt pans. Wells & Slack-Smith (1981) reported it in the higher tidal levels of the mangrove forest, including Ceriops and Aegialitis zones, on trunks up to 2 m above ground level, at densities of up to 1.65 /m 2. Remarks. As observed by Houbrick (1986), this species is closely related to C. anticipata (see Remarks on the latter). Houbrick (1986) reported that the distributions of the two just overlap in the Admiralty Gulf of northern Western Australia, where C. anticipata was said to be larger and more like C. reidi in sculpture (see Remarks on C. anticipata). However, photographs seen of twelve specimens from Port Warrender in the Admiralty Gulf (WAM S 46332, S 46081, S 46091; specimens from study of Wells 1981) all appeared to be small C. reidi. Sympatric occurrence of the two species remains to be confirmed. It is possible that closer study may reveal hybridization and intergradation in the narrow region of potential overlap. The principal differences between typical examples of the two are the larger size of C. reidi (46–61 mm, cf. 24–48 mm in C. anticipata) and its whitish shell colour with brown spiral grooves and mauve to lilac aperture (cf. brown shell with darker ridges and no mauve coloration within aperture in C. anticipata; Fig. 10 A–O). There are also subtle differences in shell sculpture; in C. anticipata the axial ribs are narrower and more prominent, stronger and more distant on the final whorl after the ventrolateral varix, and the spiral cords are usually narrower (broad and strap-like in C. reidi). The animal of C. reidi is paler in colour, but it is not known if this difference is consistent across the range of both species. Cerithidea reidi is the largest member of the genus. Its size is rivalled only by C. obtusa from Southeast Asia, but that species has a broader shell with more swollen whorls, a more heavily thickened aperture and brown cords (Fig. 8)., Published as part of Reid, David G., 2014, The genus Cerithidea Swainson, 1840 (Gastropoda: Potamididae) in the Indo-West Pacific region, pp. 1-65 in Zootaxa 3775 (1) on pages 36-37, DOI: 10.11646/zootaxa.3775.1.1, http://zenodo.org/record/285731, {"references":["Houbrick, R. S. (1986) Cerithidea reidi, spec. nov., from Western Australia. Veliger, 28, 280 - 286.","Allan, J. (1950) Australian Shells. Georgian House, Melbourne, 470 pp.","Hinton, A. G. (1978) Guide to Australian Shells. Robert Brown, Port Moresby, 82 pp.","Wells, F. E. & Bryce, C. W. (1986) Seashells of Western Australia. Western Australian Museum, Perth, 207 pp.","Lamarck, J. B. P. A. de (1822) Histoire Naturelle de Animaux sans Vertebres. Vols 6 (2), 7. Paris, 232 pp., 711 pp.","Wells, F. E. & Bryce, C. W. (1988) Seashells of Western Australia. 2 nd edition. Western Australian Museum, Perth, 207 pp.","Cecalupo, A. (2006) Elenco della famiglia Cerithiidae Ferussac, 1822 (Prosobranchia). Nomenclatura delle specie conosciute o poco note e relativa revisione sistematica. Attuali. Vol. II. Quaderni della Civica Stazione Idrobiologica di Milano, 27, 1 - 373.","Willan, R. C. (2013) A key to the potamidid snails (longbums, mudcreepers and treecreepers) of northern Australia. Northern Territory Naturalist, 24, 68 - 80.","Reid, D. G., Claremont, M., Smith, L., Shamoto, M., Glaubrecht, M. & Ozawa, T. (2013) Mosaics in the mangroves: allopatric diversification of tree-climbing mudwhelks (Gastropoda: Potamididae: Cerithidea) in the Indo-West Pacific. Biological Journal of the Linnean Society, 110, 564 - 580.","Wilson, B. R. (1993) Australian marine shells. Prosobranch gastropods. Part I. Odyssey Publishing, Kallaroo, Western Australia, 407 pp.","Wells, F. E. & Slack-Smith, S. M. (1981) Zonation of molluscs in a mangrove swamp in the Kimberley, Western Australia. In: Western Australian Museum, Biological Survey of Mitchell Plateau and Admiralty Gulf, Kimberley, Western Australia. Western Australian Museum, Perth, pp. 265 - 274."]}

Cerithidea charbonnieri (Petit de la Saussaye, 1851) (Figures 2 A, 6, 7M–V) Cerithium sp. Adams, in Adams & Reeve, 1848: 44. Cerithium charbonnieri Petit de la Saussaye, 1851: 264–265, pl. 7, fig. 7 (Borneo; 1 syntype MNHN 25689; Cecalupo 2005: pl. 30, fig. 12; Fig. 7 V). Sowerby, 1855: 886–887, pl. 186, fig. 279. Cerithidea charbonnieri — H. Adams & A. Adams, 1854: 292–293, pl. 31, fig. 2 (living animal; Cecalupo, 2006: 46, fig.). Adams, 1855: 83. Sowerby, 1866: sp. 12, pl. 2, fig. 12 a, b. Thach, 2005: 48, pl. 10, fig. 11. Reid et al., 2013: Fig. 1 (phylogeny), 2 (map). Potamides (Aphanistylus) charbonnieri — Fischer, 1884: 682, fig. 450. Crosse & Fischer, 1890: 17. Fischer, 1891: 164. Potamides (Cerithidea) charbonnieri — Tryon, 1887: 164, pl. 34, fig. 96 (as charbonieri, but correct in caption). von Martens, 1897 a: 190. Aphanistylus charbonnieri — Morlet, 1889: 144. Cerithium (Aphanistylus) charbonnieri — Kobelt, 1890 a: 47 –49, pl. 10, figs 2, 3 (as charbonieri). Cerithidea (Cerithidea) charbonnieri — Brandt, 1974: 194, pl. 14, fig. 56. Cerithidea (Aphanistylus) charbonnieri — Cecalupo, 2005: 314, pl. 30, fig. 12. Cecalupo, 2006: 46, text fig. 186 (in part, includes C. quoyii). Cerithium unicarinatum Metcalfe, 1852: 73 –74 (Borneo; 3 syntypes NHMUK 20130226, seen; Sowerby 1855: pl. 186, fig. 279; Sowerby 1866: pl. 2, fig. 12 a, b; Fig. 7 T, U; not Woodward, 1833). Cerithidea rhizophorarum — Hamli et al., 2013: 412 –418, fig. 2 b (as rizophorarum; not A. Adams, 1855) Taxonomic history. The first record of this species was a brief account of the red and yellow animal of “a very handsome Cerithium ” found in Borneo, by A. Adams (in Adams & Reeve 1848: 44; figured by H. Adams & A. Adams, 1854: pl. 31, fig. 2). Following its formal description by Petit de la Saussaye in 1851, the species has been correctly recognized by almost all subsequent authors, on account of its distinctive shell. However, only three original figures have been published since 1890 (Brandt 1974; Thach 2005; Hamli et al. 2013). Metcalfe (1852) named the species Cerithium unicarinatum, but the synonymy with C. charbonnieri was recognized shortly afterwards (Adams 1855; Sowerby 1855). Diagnosis. Shell: delicate; sutures relatively indistinct, periphery sharply angled; sinuous apertural margin, weak anterior canal, straight columella; 12–24 narrow axial ribs on penultimate whorl; ventrolateral varix is usually represented by a previous lip at about 180 °; no spiral sculpture above periphery; keel with brown line. SE Asia, Sumatra, Borneo. COI GenBank HE 680203 – HE 680206. Material examined. 28 lots. Shell (Fig. 7 M–V): H = 30.1 –46.0 mm (50 mm, Brandt 1974). Shape elongated conical (H/B = 2.41–2.93, SH = 3.03–3.61); decollate, 6–9 whorls remaining; spire whorls lightly rounded to flattened, suture relatively indistinct; spire profile straight to convex; periphery sharply angled; delicate. Adult lip only moderately flared, slightly thickened; often 1–2 previous lips on last 2 whorls; apertural margin sinuous in side view; weak anterior projection adjacent to shallow notch of canal; narrow, straight columella. Sculpture on spire of straight to slightly opisthocline axial ribs, becoming slightly curved (opisthocyrt) and weak or irregular on final 1–2 whorls, ribs narrow, interspaces 2–3 times width of ribs, 12–24 ribs on penultimate whorl; spiral sculpture absent above periphery; base with 11–13 striae, of which peripheral one forming a sharp keel. Ventrolateral varix usually developed as a previous lip at 170–210 °. Surface with faint, fine spiral microstriae on periostracum; often worn smooth (removing ribs) in old shells. Colour: pale fawn, brown line at periphery; periostracum olive brown; peripheral band visible within aperture. Animal (Fig. 2 A): Snout dark grey with yellow spots, 2 transverse black bands at base, tip red; tentacles yellowish grey with black transverse lines, black band across eye, pink base behind eye; sides of foot pinkish grey with dark grey mottling, edge red; mantle edge pale grey (red colour fading to pink in ethanol-preserved specimens). Range (Fig. 6): mainland of SE Asia, Sumatra, Borneo. Records: Malaysia: Teluk Anson, Perak (MNHN); Lubok Antu, Sarawak (NHMUK 20130236); Limbang, Sarawak (Hamli et al. 2013); Brunei: Sungai Batu Apoi Lama, Temburong Dist. (NTM P 010209). Thailand: Tapi River estuary, Surat Thani (NHMUK 20100424; AM; USNM 794070). Cambodia: Kampot (Morlet 1889). Vietnam: Long An (ANSP 330601); Nha Trang (S. Tracey). Indonesia: Boeloengan, Kalimantan (USNM 878697); Banjarmasin, Kalimantan (RMNH); Palembang, Sumatra (MNHN); Padang, Sumatra (RMNH). Both Kobelt (1890 a) and Fischer (1891) included Java in the distribution, but the species was not reported from Java by Van Benthem Jutting (1956). A record from Timor (“Coll. Dennis, 1910 ”, MNHN) is probably incorrect, because this is far outside the recorded range. The occurrence appears to be localized; in Thailand it is known from only a single estuary (Brandt 1974). Mabille (1887) recorded both C. charbonnieri and C. sinensis from Tonkin (i.e. northern Vietnam). However, specimens of the former were said to be immature and it is more likely that they were in fact juvenile C. sinensis, since C. charbonnieri has not been reliably reported from within the more northerly range of C. sinensis. Crosse & Fischer (1890) listed it as the only Cerithidea from Halong Bay, northern Vietnam, and again this is likely to refer to C. sinensis. Habitat and ecology. Common on leaf bases of Nypa palms, to 1.5 m above ground, behind narrow fringe of Avicennia trees facing estuary (Tapi River, Surat Thani, Thailand). A. Adams (in Adams & Reeve 1848: 44) reported that it was frequently found “crawling languidly on the leaves of the Pontedera [sic, i.e. Pontederia] and sedges in the fluviatile marshes on the banks of the rivers in many parts of Borneo, and many miles in the interior where the water is perfectly fresh”. It has been recorded at Sibu, Sarawak (Hamli et al. 2013), 60 km from the open sea. In Brunei it has been found on the bases of Acrostichum ferns in a riverine mangrove forest fringing a small stream (NTM P 010209). Remarks. Shells of this species are scarce in museum collections. The recorded localities are few, but it can be common where it occurs. It is used as food in Sarawak (Hamli et al. 2013). The shells of the largest and oldest animals are often severely worn (by abrasion and/or soil acidity) on the apertural side of the spire. The combination of sharp peripheral keel with solely axial sculpture makes this species easily recognized. Two genus-level taxa have been based on this species but there is no apparent morphological or phylogenetic justification for this (see Taxonomic History of the genus Cerithidea). It forms a well-supported clade with C. balteata and C. sinensis in molecular phylogenies based on COI and 28 S (Reid et al. 2013; Fig. 1), but which of these is its sister species is not clear., Published as part of Reid, David G., 2014, The genus Cerithidea Swainson, 1840 (Gastropoda: Potamididae) in the Indo-West Pacific region, pp. 1-65 in Zootaxa 3775 (1) on pages 25-27, DOI: 10.11646/zootaxa.3775.1.1, http://zenodo.org/record/285731, {"references":["Petit de la Saussaye, M. S. (1851) Description de coquilles nouvelles. Journal de Conchyliologie, 2, 259 - 269.","Adams, A. & Reeve, L. (1848 - 1850) Mollusca. In: Adams, A. (Ed.), The Zoology of the Voyage of H. M. S. Samarang. Reeve, Benham & Reeve, London, x + 87 pp.","Cecalupo, A. (2005) Elenco della famiglia Cerithiidae Ferussac, 1822 (Prosobranchia). Nomenclatura delle specie conosciute o poco note e relativa revisione sistematica. Fossili e Attuali. Vol. I. Quaderni della Civica Stazione Idrobiologica di Milano, 26, 1 - 366.","Sowerby, G. B. II (1855) Monograph of the genus Cerithium, Adanson. In: Sowerby, G. B. II. (Ed.), Thesaurus Conchyliorum, or Figures and Descriptions of Recent Shells. Vol. 2. part 16. Sowerby, London, pp. 847 - 899.","Cecalupo, A. (2006) Elenco della famiglia Cerithiidae Ferussac, 1822 (Prosobranchia). Nomenclatura delle specie conosciute o poco note e relativa revisione sistematica. Attuali. Vol. II. Quaderni della Civica Stazione Idrobiologica di Milano, 27, 1 - 373.","Adams, A. (1855) A monograph of Cerithidea, a genus of Mollusca, with descriptions of several new species, from the collection of High Cuming, Esq,: to which are added, descriptions of two new species of Colina, and one of Donax. Proceedings of the Zoological Society of London, 1854, 83 - 87. http: // dx. doi. org / 10.1111 / j. 1469 - 7998.1854. tb 07236. x","Sowerby, G. B. II (1866) Monograph of the genus Cerithidea. In: Reeve, L. A. (Ed.), Conchologia Iconica. Vol. 15. L. A. Reeve, London, pls 1 - 4.","Thach, N. N. (2005) Shells of Vietnam. Conchbooks, Hackenheim, 338 pp.","Reid, D. G., Claremont, M., Smith, L., Shamoto, M., Glaubrecht, M. & Ozawa, T. (2013) Mosaics in the mangroves: allopatric diversification of tree-climbing mudwhelks (Gastropoda: Potamididae: Cerithidea) in the Indo-West Pacific. Biological Journal of the Linnean Society, 110, 564 - 580.","Crosse, H. & Fischer, P. (1890) Mollusques marins de la Baie d'Halong (Tonkin). Journal de Conchyliologie, 38, 14 - 19.","Fischer, P. (1891) Catalogue et distribution geographique des mollusques terrestres, fluviatiles et marins d'une partie de l'Indo- Chine (Siam, Laos, Cambodge, Annam, Tonkin). Bulletin de la Societe d'Histoire Naturelle d'Autun, 4, 84 - 276.","Tryon, G. W. (1887) Manual of Conchology. Vol. 9. Tryon, Philadelphia, 488 pp.","Martens, E. von (1897 a) Suss- und Brackwasser-Mollusken des Indischen Archipels. In: Weber, M. (Ed.), Zoologische Ergebnisse einer Reise in Niederlandisch Ost-Indien, Vol. 4. E. J. Brill, Leiden, pp. 1 - 331.","Morlet, L. (1889) Catalogue des coquilles recueillis, par M. Pavie, dans le Cambodge et le Royaume de Siam, et description d'especes nouvelles. Journal de Conchyliologie, 37, 121 - 199.","Kobelt, W. (1890 b) Eine neue Cerithidea. Nachrichtsblatt der deutschen Malakozoologischen Gesellschaft, 1890, 60 - 61.","Brandt, R. A. M. (1974) The non-marine aquatic Mollusca of Thailand. Archiv fur Molluskenkunde, 105, 1 - 423.","Metcalfe, W. (1852) An enumeration of species of recent shells, received by W. J. Hamilton, Esq., from Borneo, in November 1850, with descriptions of the new species. Proceedings of the Zoological Society of London, 1851, 70 - 74. http: // dx. doi. org / 10.1111 / j. 1096 - 3642.1851. tb 01132. x","Woodward, S. P. (1833) An Outline of the Geology of Norfolk. John Stacy, Norwich, 54 pp.","Hamli, H., Idris, M. H., Abu Hena, M. K., Wong, S. K. & Arshad, A. (2013) Checklist and habitat of edible gastropods from Sarawak, Malaysia. Journal of Fisheries and Aquatic Sciences, 8, 412 - 418. http: // dx. doi. org / 10.3923 / jfas. 2013.412.418","Van Benthem Jutting, W. S. S. (1956) Systematic studies on the non-marine Mollusca of the Indo-Australian archipelago. V. Critical revision of the Javanese freshwater gastropods. Treubia, 23, 259 - 477.","Mabille, J. (1887) Sur quelques mollusques du Tonkin. Bulletins de la Societe Malacologique de France, 4, 73 - 164."]}

Cerithidea houbricki new species (Figures 3 N–R, 4) Cerithidea rhizophorarum — Cernohorsky, 1972: 61, pl. 13, fig. 3 (as rhizoporarum; not A. Adams, 1855). Poraituk & Ulijaszek, 1981: 13, pl. 2 b (as rhizoporarum; not A. Adams, 1855). Cerithidea n. sp. A Reid et al., 2013: fig. 2 (map). Types. Holotype AM C. 126895 (Fig. 3 O–Q) and five paratypes AM C. 478233 (Fig. 3 N), Koravake, Gulf Prov., Papua New Guinea. Etymology: Richard S. Houbrick was the pioneer of the modern anatomical and phylogenetic study of the Cerithioidea, including the genus Cerithidea (e.g. Houbrick 1984, 1986, 1988). He was a mentor for me and other young malacologists until his untimely death in 1993 (Harasewych & Kabat 1995). He described C. reidi in 1986 and, after 28 years, I am delighted to return his compliment. Taxonomic history. A shell of this species (Fig. 3 R) was illustrated by Cernohorsky (1972) as C. rhizophorarum; this misidentification was repeated by Poraituk & Ulijaszek (1981), whose material was identified by P. Colman of the AM, where it is now deposited and is the type collection of the new species. The existence of this new species was noted by Reid et al. (2013). Diagnosis. Shell: large, solid, elongated-pupoidal with expanded and thickened aperture, periphery rounded; 19–29 rounded axial ribs (remaining strong and becoming rugose on final whorl); ventrolateral varix weak, at 220– 260 °; 4 spiral ridges above periphery on final 1–2 whorls; banded pattern. New Guinea. Material examined. 2 lots. Shell (Fig. 3 N–R): H = 29.9 –41.0 mm. Shape elongated-pupoidal (H/B = 2.16–2.40, SH = 2.85–3.04); decollate, 6–7 whorls remaining; spire whorls lightly rounded; spire profile convex; periphery rounded to slightly angled; solid. Adult lip thickened, moderately flared; apertural margin planar in side view; weak anterior projection adjacent to canal. Sculpture on spire of straight to slightly opisthocline or curved (opisthocyrt) axial ribs, occasionally bifurcating posteriorly (adapically), remaining strong and becoming rugose on final whorl, usually continuing as rugose ribs on base, ribs rounded, interspaces 1–1.5 width of ribs, 19–29 ribs on penultimate whorl; spiral sculpture of 4 strong cords above periphery on final 1–2 whorls, producing nodules at intersection with axial ribs; base with 4–7 cords. Ventrolateral varix weakly developed at 220–260 °. Surface with faint, fine spiral microstriae on periostracum, satin sheen. Colour: fawn, broad brown band above aperture, spiral ridges brownish, brown band adjacent to columella; occasionally entirely dark brown; aperture brown. Animal: Unknown. Range (Fig. 4): New Guinea. Records: Papua New Guinea: Koravake, Gulf Prov. (AM C. 126895). Indonesia: Samberbaba, Yapen I., East Papua (USNM 835667). The specimen from Yapen is doubtfully assigned to this species (see below). Habitat and ecology. Poraituk & Ulijaszek (1981) reported it to be common in the upper reaches of the Pie River estuary, 20–30 km from the open sea, at the extreme landward fringe of the mangrove forest. It occurred in clusters on the bases of tree trunks and on mud and rotten wood. Other molluscs present were Polymesoda and Neritina, so salinity was evidently low. Remarks. Only two samples of this species were available, the six type specimens and a single specimen from Yapen, all dry shells. The relatively solid shell, slightly pupoidal outline (in the type collection), weak ventrolateral varix and axial ribs continuing strongly beyond the varix, all indicate that its closest relationship is likely to be with C. weyersi. The new species is distinguished from this likely sister by larger size (to 41 mm, cf. 33 mm) and by the 4 spiral ridges above the periphery on the body whorl, producing a nodulose or rugose sculpture. The island of Yapen is geographically closer to the nearest record of C. weyersi from Halmahera, than it is to the other locality of C. houbricki in the Gulf of Papua (Fig. 4), but the shell from Yapen is not similar to the typical C. weyersi from Halmahera. Compared with the type collection, the specimen from Yapen is narrower, more cylindrical in shape, smaller (H = 34.7 mm) and the axial ribs do not continue on to the base (Fig. 3 R; the same specimen was figured by Cernohorsky, 1972), although these differences are no greater than those seen among specimens of C. weyersi (compare Fig. 3 A–M). A recent expedition to Madang, northern New Guinea, failed to find any Cerithidea species in mangrove habitats (P. Bouchet, pers. comm.) and there is little development of mangrove forests along the entire north coast of New Guinea. Mangroves line most of the south coast of New Guinea and collections of Cerithidea species are available from several localities (see C. anticipata and C. balteata), but the present species has not been recorded there. Phylogeographic studies of marine invertebrates have suggested that Cenderawasih Bay (including the island of Yapen) is isolated by marine currents (Barber et al. 2006; Carpenter et al. 2011). For these reasons it is unlikely that the collections from Yapen and Koravake are conspecific, but more material is needed to investigate this further. Like its probable sister, C. weyersi, this species may have remained poorly known and rarely collected because of its transitional habitat between marine mangrove forests and freshwater streams. Further study of additional material, and application of molecular techniques, is very desirable. Poraituk & Ulijaszek (1981) reported that this species was a significant component of the diet of indigenous people of the Purari Delta, Papua New Guinea., Published as part of Reid, David G., 2014, The genus Cerithidea Swainson, 1840 (Gastropoda: Potamididae) in the Indo-West Pacific region, pp. 1-65 in Zootaxa 3775 (1) on pages 17-18, DOI: 10.11646/zootaxa.3775.1.1, http://zenodo.org/record/285731, {"references":["Cernohorsky, W. O. (1972) Marine Shells of the Pacific. Vol. 2. Pacific Publications, Sydney, 411 pp.","Adams, A. (1855) A monograph of Cerithidea, a genus of Mollusca, with descriptions of several new species, from the collection of High Cuming, Esq,: to which are added, descriptions of two new species of Colina, and one of Donax. Proceedings of the Zoological Society of London, 1854, 83 - 87. http: // dx. doi. org / 10.1111 / j. 1469 - 7998.1854. tb 07236. x","Poraituk, S. P. & Ulijaszek, S. (1981) Molluscs in the subsistence diet of some Purari Delta people. Purari River (Wabo) Hydroelectric Scheme Environmental Studies, 20, 1 - 30.","Reid, D. G., Claremont, M., Smith, L., Shamoto, M., Glaubrecht, M. & Ozawa, T. (2013) Mosaics in the mangroves: allopatric diversification of tree-climbing mudwhelks (Gastropoda: Potamididae: Cerithidea) in the Indo-West Pacific. Biological Journal of the Linnean Society, 110, 564 - 580.","Houbrick, R. S. (1984) Revision of higher taxa in genus Cerithidea (Mesogastropoda: Potamididae) based on comparative morphology and biological data. American Malacological Bulletin, 2, 1 - 20.","Houbrick, R. S. (1986) Cerithidea reidi, spec. nov., from Western Australia. Veliger, 28, 280 - 286.","Houbrick, R. S. (1988) Cerithioidean phylogeny. Malacological Review, Suppl. 4, 88 - 128.","Harasewych, M. G. & Kabat, A. R. (1995) Richard S. Houbrick (1937 - 1993): biographical sketch and malacological contributions. Nautilus, 108, 61 - 66.","Barber, P. H., Erdmann, M. V. & Palumbi, S. R. (2006) Comparative phylogeography of three codistributed stomatopods: origins and timing of regional lineage diversification in the Coral Triangle. Evolution, 60, 1825 - 1839. http: // dx. doi. org / 10.1111 / j. 0014 - 3820.2006. tb 00526. x","Carpenter, K. E., Barber, P. H., Crandall, E. D., Ablan-Lagman, M. C. A., Ambariyanto, Mahardika, G. N., Manjaji-Matsumoto, B. M., Juinio-Menez, M. A., Santos, M. D., Starger, C. J. & Toha, A. H. A. (2011) Comparative phylogeography of the Coral Triangle and implications for marine management. Journal of Marine Biology, 2011, Article ID 396982, 1 - 14. http: // dx. doi. org / 10.1155 / 2011 / 396982"]}

Cerithidea weyersi Dautzenberg, 1899 (Figures 2 H, 3 A–M, 4) Cerithidea ornata — Boettger, 1890 a: 167 (synonymy fide Van Benthem Jutting 1956; not Sowerby, 1855 = C. balteata). Potamides (Cerithidea) ornatus —von Martens, 1897 a: 189 (not Sowerby, 1855; includes C. balteata). Potamides ornatus — Leschke, 1914: 437 (not Sowerby, 1855). Cerithidea (Aphanistylus) weyersi Dautzenberg, 1899: 8 –10, pl. 2, fig. 1, 1a, 1 b (delta de la rivière d’Indrapoera, Sumatra [Indrapura, Indonesia]; 3 syntypes IRSNB MT 2772 (Dautzenberg, 1899: pl. 2, fig. 1, 1a, 1 b; Cecalupo 2006: 170, fig.; Fig. 3 A); 3 syntypes IRSNB MT 2179; 54 syntypes IRSNB; 55 + 36 probable syntypes IRSNB; 5 syntypes RMNH MOL. 175859). Cerithidea weyersi — Van Benthem Jutting, 1956: 436 –438, fig. 105. Dharma, 2005: 92, pl. 21, fig. 7 a, b. Mujiono, 2009: 53 – 54, fig. 1 F. Reid et al., 2013: figs 1 (shell, phylogeny), 2 (map). Cerithidea (Cerithidea) weyersi — Brandt, 1974: 193 –194, pl. 14, figs 54, 55. Cerithidea (Cerithideopsis) scalariformis — Cecalupo, 2005: 320, pl. 33, fig. 8 (not Say, 1825). Cerithidea rhizophorarum — Cecalupo, 2006: 170, 234, 256, fig. (not A. Adams, 1855). Cerithidea obtusa — Ahmed & Mahmoud, 2007: 10, 95, fig. p. 10 (in part, includes C. obtusa; not Lamarck, 1822). Cerithidea quadrata — Ahmed & Mahmoud, 2007: 96, fig. (not Sowerby, 1866). Taxonomic history. This species is poorly known and has been widely misidentified. The first misidentification was by Boettger (1890 a), whose record of ‘ C. ornata ’ was based on a specimen of C. weyersi from Java, according to Van Benthem Jutting (1956). The reference by von Martens (1897 a) is included in the synonymy only because he listed Boettger’s (1890 a) record of ‘ C. ornata ’ from Java; the rest of his text applies entirely to C. balteata (his record of ‘ C. ornata ’ from ‘Sumatra’ – where only C. weyersi occurs – is a misquotation of ‘Soemba’ [Sumba] – based on 10 specimens of C. balteata collected by Ten Kate, in RMNH – in Schepman 1892; see Range, below). Following its naming by Dautzenberg (1899), the identity of the species was correctly established by Van Benthem Jutting (1956) and Brandt (1974). Diagnosis. Shell: small, solid, elongated-pupoidal with expanded aperture, periphery rounded; 18–30 rounded axial ribs (remaining strong throughout final whorl); ventrolateral varix absent or weak, at 230–270 °; no spiral sculpture above periphery; often a strong pattern of spiral lines. Eastern Indian Ocean, Java, Borneo, Moluccas. COI GenBank HE 680298. Material examined. 26 lots. Shell (Fig. 3 A–M): H = 15.8–33 mm. Shape elongated-pupoidal (H/B = 1.91–2.43, SH = 2.87 –3.00); decollate, 5–7 whorls remaining; spire whorls slightly rounded; spire profile convex, concave at attenuated apex if only partially decollate (Fig. 3 H, K); periphery rounded; relatively solid. Adult lip moderately flared and thickened, sometimes greatly thickened (e.g. 6.2 mm thick in shell H = 24.6 mm; Fig. 3 M); rarely 1–2 previous lips on last whorl; apertural margin planar in side view; weak anterior projection adjacent to canal. Sculpture on spire of straight to slightly opisthocline axial ribs, becoming slightly curved (opisthocyrt) on final 1–3 whorls, remaining strong on final whorl and continuing weakly on base, ribs rounded, ribs and interspaces of equal width, 18–30 ribs on penultimate whorl; spiral sculpture absent above periphery; base with 9–11 indistinct striae. Ventrolateral varix absent or weakly developed at 230–270 °. Surface with faint, fine spiral microstriae on periostracum, satin sheen. Colour: fawn, usually a brown band just below periphery and brown line adjacent to columella, usually 3–4 brown lines above periphery; occasionally without lines (Fig. 3 J) or lines may fuse so shell is entirely brown above periphery (Fig. 3 I); periostracum pale to dark brown; bands visible within aperture. Animal (Fig. 2 H): Fawn, mottled with brown; 3 brown to black bands across snout, becoming darker distally; tentacles with transverse black lines, black band across eye; sides of foot mottled with grey, brown and orange brown; mantle edge fawn. Range (Fig. 4): Bangladesh, W Thailand, W Sumatra, Java, Borneo, Moluccas. Records: Bangladesh: Char Kukrimukri, Bhola (NHMUK 20130231); R. Ganges (ZMB). Thailand: Ban Nai Sa, Krabi Prov. (RMNH); Kantang, Trang Prov. (NHMUK 20130230; RMNH; AM; USNM 777194; ANSP 330948). Indonesia: Mempawah, Kalimantan (ZMB); Marang, Sumatra (ANSP 72450); Indrapoera, Sumatra (RMNH); Welkomst Bay, Banten, Java (USNM 261761); Anyer–Labuhan road, Java (ZMB 106.427); Tji Mandiri estuary, Java (RMNH); Pangandaran– Cipatujah road, Java (ZMB 106.428); S Madura, Java (RMNH); W Gumbrih, Bali (ZMB 191.521); Ternate, Halmahera (RMNH 175855). The records are mostly from the western and southern edge of continental Southeast Asia and the former Sundaland, but with two exceptions (listed above). One, from western Borneo, is a single shell collected at Mempawah by von Martens in the late nineteenth century and is considered reliable. The other is from Halmahera, based on eleven specimens collected in 1914 by E.F. Jochim. The identification is not in question but, since there are no records between this locality and Bali, the occurrence requires confirmation. In addition to the known range in Thailand, Sumatra and Java, Brandt (1974) included Buru, Sumba and Timor in eastern Indonesia, and Luzon and Negros in the Philippines, in the distribution of this species. These last five records were listed without authority and appear to have been taken from an identical list of localities given by von Martens (1897 a) for C. ornata [= C. balteata]. In this, Brandt appears to have followed Van Benthem Jutting’s (1956) inclusion of von Martens’ ‘ C. ornata ’ in the synonymy of C. weyersi. Nevertheless, von Martens’ text clearly applies largely to C. balteata (see Taxonomic History, above). Therefore, the only known record of C. weyersi from the Banda or Molucca Seas is that from Halmahera, mentioned above. Habitat and ecology. Habitats recorded on museum labels include: “Nipa-palm swamp” in Trang Prov., Thailand (leg. R.A.M. Brandt, NHMUK 20130230); “river 200 m from sea” in Java (leg. T. von Rintelen, ZMB); “bases of Sonneratia trees in mangrove plantation” in Bangladesh (leg. L.M. Cook, NHMUK 20130231). At Gumbrih, Bali, it was collected on the banks of a short lowland river, at a site with brackish (but seasonally fresh) water, together with freshwater cerithioideans (Stenomelania, Tarebia, ‘ Thiara ’ scabra) (M. Glaubrecht, pers. comm.). In Java, Van Benthem Jutting (1956) recorded C. weyersi from river estuaries that were brackish at high tide and fresh at low tide, while Mujiono (2009) reported it on roots and trunks of mangroves. In Bangladesh it has been said to climb on trees, but a figure shows it attached to grass (Ahmed & Mahmoud 2007: 10, 96). In Krabi Prov., Thailand, it was found on vegetation at the margin of a fast-flowing river, shaded by mangrove trees, at the upstream limit of brackish influence (B. Craig, pers. comm.). Remarks. This is a little known species, poorly represented in museum collections. Its habitat is apparently towards the freshwater end of the estuarine range and occasional large samples suggest that it can sometimes be abundant. Possibly, this transitional habitat might explain why it has seldom been collected, since it does not occur in typically marine mangrove environments or in fully freshwater streams. Its distribution is peculiar. The majority of the records are from the eastern margin of the Indian Ocean (Bangladesh, W Thailand, Sumatra, Java and Bali), an area that corresponds with the margin of Sundaland during glacial low sea-level stands (Voris 2000). This implies that the distribution could be relictual, because of limited ability to recolonize the shelf when sea levels rose in the Holocene. However, the records from Borneo and Halmahera do not support this. The recorded distribution is likely to be incomplete. In a molecular phylogeny based only on mitochondrial COI sequences (Reid et al. 2013; Fig. 1), the single individual of C. weyersi forms one of three basal branches in the genus. Morphological comparison suggests that it is sister to C. houbricki, and their distinction is discussed in the Remarks on that species., Published as part of Reid, David G., 2014, The genus Cerithidea Swainson, 1840 (Gastropoda: Potamididae) in the Indo-West Pacific region, pp. 1-65 in Zootaxa 3775 (1) on pages 13-17, DOI: 10.11646/zootaxa.3775.1.1, http://zenodo.org/record/285731, {"references":["Dautzenberg, P. (1899) Contribution a la faune malacologique de Sumatra (recoltes de M. J. - L. Weyers). Annales de la Societe Royale Malacologique de Belgique, 34, 1 - 26.","Boettger, O. (1890 a) Ad. Strubell's Konchylien aus Java I. Bericht uber die Senckenbergische naturforschende Gesellschaft in Frankfurt am Main, 1890, 137 - 173.","Van Benthem Jutting, W. S. S. (1956) Systematic studies on the non-marine Mollusca of the Indo-Australian archipelago. V. Critical revision of the Javanese freshwater gastropods. Treubia, 23, 259 - 477.","Sowerby, G. B. II (1855) Monograph of the genus Cerithium, Adanson. In: Sowerby, G. B. II. (Ed.), Thesaurus Conchyliorum, or Figures and Descriptions of Recent Shells. Vol. 2. part 16. Sowerby, London, pp. 847 - 899.","Martens, E. von (1897 a) Suss- und Brackwasser-Mollusken des Indischen Archipels. In: Weber, M. (Ed.), Zoologische Ergebnisse einer Reise in Niederlandisch Ost-Indien, Vol. 4. E. J. Brill, Leiden, pp. 1 - 331.","Leschke, M. (1914) Zur Molluskenfauna von Java und Celebes. Mitteilungen aus dem Naturhistorischen Museum in Hamburg, 31, 205 - 284.","Cecalupo, A. (2006) Elenco della famiglia Cerithiidae Ferussac, 1822 (Prosobranchia). Nomenclatura delle specie conosciute o poco note e relativa revisione sistematica. Attuali. Vol. II. Quaderni della Civica Stazione Idrobiologica di Milano, 27, 1 - 373.","Dharma, B. (2005) Recent & Fossil Indonesian Shells. ConchBooks, Hackenheim, 424 pp.","Mujiono, N. (2009) Mudwhelks (Gastropoda: Potamididae) from mangroves of Ujung Kulong National Park, Banten. Jurnal Biologi, 13, 51 - 56.","Reid, D. G., Claremont, M., Smith, L., Shamoto, M., Glaubrecht, M. & Ozawa, T. (2013) Mosaics in the mangroves: allopatric diversification of tree-climbing mudwhelks (Gastropoda: Potamididae: Cerithidea) in the Indo-West Pacific. Biological Journal of the Linnean Society, 110, 564 - 580.","Brandt, R. A. M. (1974) The non-marine aquatic Mollusca of Thailand. Archiv fur Molluskenkunde, 105, 1 - 423.","Cecalupo, A. (2005) Elenco della famiglia Cerithiidae Ferussac, 1822 (Prosobranchia). Nomenclatura delle specie conosciute o poco note e relativa revisione sistematica. Fossili e Attuali. Vol. I. Quaderni della Civica Stazione Idrobiologica di Milano, 26, 1 - 366.","Say, T. (1825) Descriptions of some new species of fresh water and land snails of the United States. Journal of the Academy of Natural Sciences of Philadelphia, 5, 119 - 131.","Adams, A. (1855) A monograph of Cerithidea, a genus of Mollusca, with descriptions of several new species, from the collection of High Cuming, Esq,: to which are added, descriptions of two new species of Colina, and one of Donax. Proceedings of the Zoological Society of London, 1854, 83 - 87. http: // dx. doi. org / 10.1111 / j. 1469 - 7998.1854. tb 07236. x","Ahmed, A. T. A. & Mahmoud, M. M. (2007) Family Potamididae. In: Siddiqui, K. U., Islam, M. A., Kabir, S. M. H., Ahmad, M., Ahmed, A. T. A., Rahman, A. K. A., Haque, E. U., Ahmed, Z. U., Begum, Z. N. T., Hassan, M. A., Khondker, M. & Rahman, M. M. (Eds.), Encyclopedia of Flora and Fauna of Bangladesh. Vol. 17. Molluscs. Asiatic Society of Bangladesh, Dhaka, pp. 9, 10, 93 - 98.","Lamarck, J. B. P. A. de (1822) Histoire Naturelle de Animaux sans Vertebres. Vols 6 (2), 7. Paris, 232 pp., 711 pp.","Sowerby, G. B. II (1866) Monograph of the genus Cerithidea. In: Reeve, L. A. (Ed.), Conchologia Iconica. Vol. 15. L. A. Reeve, London, pls 1 - 4.","Schepman, M. M. (1892) Note XXVIII. Land- and freshwater shells collected by Dr H. Ten Kate in Soemba, Timor and other East-Indian islands. Notes from the Leyden Museum, 14, 145 - 160.","Voris, H. K. (2000) Maps of Pleistocene sea levels in Southeast Asia: shorelines, river systems and time durations. Journal of Biogeography, 27, 1153 - 1167. http: // dx. doi. org / 10.1046 / j. 1365 - 2699.2000.00489. x"]}

Cerithidea balteata A. Adams, 1855 (Figures 5, 6) Cerithidea balteata A. Adams, 1855: 84 (Island of Ticao [Philippines]; lectotype here designated NHMUK 20130222 / 1, Fig. 5 A, 2 paralectotypes NHMUK 20130222 /2, 3). Hidalgo, 1904 –1905: 202–203. Reid et al., 2013: figs 1 (shell, phylogeny), 2 (map). Cerithium (Cerithidea) balteatum — Kobelt, 1895: 177, pl. 33, figs 5, 6 (Cecalupo 2005: pl. 31, fig. 11). Cerithidea balteata —Casto de Elera, 1896: 329. Cerithidea raricostata A. Adams, 1855: 85 (“Catbalonga, island of Samaar” here corrected to Calbayog, Samar [Philippines], as per original label; 3 syntypes NHMUK 20130223, seen). Cerithium (balteata var.?) rarecostatum — Kobelt, 1895: 178–179, pl. 33, figs 7, 8 (Cecalupo 2005: pl. 31, fig. 12). Cerithidea (Aphanistylus) raricostata —Casto de Elera, 1896: 330. Cerithidea raricosta — Hidalgo, 1904 –1905: 203. Cerithidea cornea A. Adams, 1855: 85 (“Borneo” here corrected to Guimaras [Philippines], as per original label; 3 syntypes NHMUK 20130224, Fig. 5 D, seen). Sowerby, 1866: sp. 21, pl. 3, fig. 21. Casto de Elera, 1896: 328. Cerithium corneum — Sowerby, 1855: 887, pl. 186, fig. 275. Potamides (Cerithidea) corneus — Tryon, 1887: 162, pl. 32, fig. 55 (as cornea). von Martens, 1897 a: 190. Cerithium (Cerithidea) corneum — Kobelt, 1890 a: 50 –51, pl. 10, figs 6, 7 (Cecalupo 2005: pl. 31, fig. 5). Cerithium ornatum ‘A. Adams’ G.B. Sowerby II, 1855: 887, pl. 186, figs 277, 278 (“ Philippines ”, here corrected to Ticao, Philippines, as per original label; lectotype here designated NHMUK 20130222 / 1, Fig. 5 A, Sowerby, 1855: pl. 186, fig. 278, 2 paralectotypes NHMUK 20130222 /2, 3; 2 paralectotypes NHMUK 20130225, Sowerby, 1855: pl. 186, fig. 277). Cerithidea ornata — Sowerby, 1866: sp. 22, pl. 4, fig. 22 a, b. Casto de Elera, 1896: 328. Hidalgo, 1904 –1905: 203. Schepman, 1909: 168 –169. Oostingh, 1925: 50. Lozouet, 2008: 284, pl. 87, figs 7, 8. Lozouet & Plaziat, 2008: 57, 112, pl. 20, figs 7– 9. Reid et al., 2008: 680 –699, figs 1, 2 (phylogeny) (in part, includes C. tonkiniana). Potamides (Cerithidea) ornata — Tryon, 1887: 162, pl. 32, fig. 64. von Martens, 1897 a: 189 (in part, includes C. weyersi fide Van Benthem Jutting 1956; as ornatus). Cerithium (Cerithidea) ornatum — Kobelt, 1890 a: 51 –52, pl. 11, figs 1, 2 (Cecalupo 2005: pl. 31, fig. 8). Cerithium (Cerithidea) quadrasi Kobelt, 1895: 176, pl. 33, figs 3, 4 (Marinduque, Philippines; 2 syntypes SMF 228128, photograph seen, 228129; Cecalupo 2005: pl. 31, fig. 9). Cerithium (Cerithidea) balteatum var. mindorensis Kobelt, 1895: 179, pl. 33, fig. 9 (Mindoro [Philippines]; holotype SMF 228127, photograph seen; Cecalupo 2005: pl. 32, fig. 28). Cerithidea (Cerithideopsis) scalariformis — Cecalupo, 2005: 316–318, pl. 31, figs 5, 9, 11, 12, pl. 32, fig. 28 (not Say, 1825; in part, includes Cerithideopsis scalariformis). Cecalupo, 2006: 34, 53, 102, 133, 179, 190, 218, 233, 236, 237 (not Say, 1825; in part, includes Cerithideopsis scalariformis). Cerithidea anticipata — Cecalupo, 2005: 316, pl. 31, fig. 8 (not Iredale, 1929; in part, includes C. anticipata). Cecalupo, 2006: 118, 176, 224 (not Iredale, 1929; in part, includes C. anticipata, Cerithideopsilla conica, Cerithideopsis largillierti). Taxonomic history. This species is poorly known and variable in appearance, accounting for its complex history. Four of the six available names were introduced in 1855 (C. balteata, C. raricostata and C. cornea by Adams; Cerithium ornatum by Sowerby). None is well known, but Cerithium ornatum has been used a little more frequently. However, Cerithium ornatum has far more often been used in an incorrect sense (for C. tonkiniana). As first reviser, I choose C. balteata for the present species, to minimise confusion (this name was also used recently for this species by Reid et al. 2013). Adams’ paper was published on 11 April 1855 and although the exact date of Sowerby’s work is not known, it can be assumed to have been later (Petit 2009) because he uses most of Adams’ new names and copied some of his descriptions. (In fact Sowerby 1855: 848 mentions only the use of a manuscript by Adams and therefore may not have seen Adams’ published monograph of Cerithidea). Sowerby did not, however, mention C. balteata. Instead he introduced Cerithium ornatum and attributed the name to Adams. One of Sowerby’s figured specimens (Sowerby 1855: pl. 186, fig. 278; here designated lectotype of Cerithium ornatum) is part of the type series of C. balteata. Sowerby assigned most cerithioideans to the genus Cerithium, so Adams’ species would have become a junior homonym of Cerithium balteatum Philippi, 1848. While this was probably the reason why Sowerby introduced the name Cerithium ornatum, he did not explicitly do so as a replacement for C. balteata. Furthermore, the description and type locality are not exactly the same as those of C. balteata (unlike, for example, Sowerby’s text for Adams’ species C. cornea, which is an exact copy of Adams 1855), and the type material includes additional specimens. For these reasons Cerithium ornatum is regarded as a new species, not a replacement name. To remove any ambiguity, the same specimen is here designated lectotype of each, so that the names become objective synonyms. Authorship of Cerithium ornatum is here attributed to Sowerby (not ‘Adams in Sowerby’ as per Petit 2009: 34, 147), because of the discrepancies between the descriptions of C. balteata and Cerithium ornatum, and because the renaming of Adams’ species was only necessary because of its classification in the broad genus Cerithium as advocated by Sowerby, whereas Adams himself used Cerithidea (Adams 1855). The type locality of C. cornea was given as “Borneo, mouths of rivers” (Adams, 1855), and this was repeated by Sowerby (1855, 1866). However, the original label accompanying the syntypes, written by the collector Hugh Cuming, reads “Isle of Guimaras in marshy places overflowed by the sea at times”. There is no firm evidence that this species occurs on the mainland of Borneo, so the type locality is here corrected. The number of synonyms reflects the variability of this species in colour, thickness and number of axial ribs. Cerithidea quadrasi appears to be a form with strong spiral lirae. A species from the Far East, C. tonkiniana, has been widely confused with the present species, under the name C. ornata. Cecalupo (2005, 2006) synonymized Cerithium ornatum with C. anticipata, and all the other synonyms listed above with the western Atlantic species Cerithideopsis scalariformis. Diagnosis. Shell: periphery rounded or slightly angled; aperture thickened and flared; 8–31 rounded axial ribs on penultimate whorl, 0–6 weaker ribs after ventrolateral varix; ventrolateral varix a strong or inconspicuous rib at (180) 210–280 °; no spiral sculpture on spire but 1–6 weak ridges above periphery on last whorl; often strongly banded or lined pattern. Philippines, Moluccas, New Guinea, Solomon Is. COI GenBank AM 932830, HE 680208, HE 680225 – HE 680228. Material examined. 111 lots. Shell (Fig. 5): H = 16.0– 37.3 mm. Shape elongated conical (H/B = 1.91–2.37, SH = 2.46–3.60); decollate, 5– 8 whorls remaining; spire whorls rounded to flattened, suture distinct; spire profile straight to slightly convex; periphery rounded to slightly angled; thin to moderate thickness. Adult lip flared, moderately thickened; occasionally a previous lip on final whorl; apertural margin planar in side view; moderate anterior projection adjacent to canal. Sculpture on spire of straight to slightly opisthocline axial ribs, becoming slightly curved (opisthocyrt) on final 1–2 whorls, ribs usually prominent, rounded, interspaces 1–2 times width of ribs, 8–31 ribs on penultimate whorl; axial ribs after ventrolateral varix usually weaker and more distant than on spire, or irregularly rugose, numbering 0–6; spiral sculpture absent on spire, but 1–6 weak ridges often develop above periphery on final whorl (rarely, ridges conspicuous between ribs on last 2–4 whorls; Fig. 4 Q–U); base with 5–10 striae, of which peripheral one may form weak keel. Ventrolateral varix usually a thickened rib at (180) 210–280 °, but may be inconspicuous or absent. Surface with faint, fine spiral microstriae on periostracum; ribs shiny when worn. Colour: fawn, usually 1–6 dark brown bands or lines above periphery and 2 on base, sometimes dark brown with indistinct pale bands; bands show through in aperture. Animal: Head grey with pink and pale yellow spots; snout grey with transverse black lines and 3 black bands, tip yellow; tentacles grey with black transverse lines, black band across eye, pink base behind eye; sides of foot pinkish grey with dark grey mottling; mantle grey, edge cream (based on ethanol-preserved specimens from Sulawesi). Range (Fig. 6): Philippines, Sulawesi, Moluccas, Lesser Sunda Is, New Guinea, Solomon Is. Records: Philippines: Port San Vicente, Palaui I., Luzon (USNM 259402); Malinao, Albay, Luzon (USNM 1165); Malubatglubut I., Palawan (USNM 233037); Port Dupon, Leyte (USNM 232971); Jordan R., Guimaras (USNM 243887); Port Tataan, Tataan I. (USNM 862783); Surigao, Mindanao (USNM 544065); Davao, Mindanao (USNM 233133). Indonesia: Manado, Sulawesi (NHMUK); Luwuk, Sulawesi (ZMB 191.377); Tembecha, Kendari, Sulawesi (ZMB 191.376); Lovina, Bali (ZMB 106.180); Waiara, Maumere, Flores (ZMB 106.573); Kupang, Timor (ZMB); Bacan, Moluccas (RMNH); Obi, Moluccas (NHMUK 1903.4.863; AM C. 33594; RMNH); Aru Is (NHMUK 1858.3.17.29); Warir I., E of Salawati I., East Papua (RMNH); Mimika R., Kokenau, East Papua (AM C. 116356); Sorendidori, E side Soepiori I., Schouten Is, East Papua (ANSP 207473). Solomon Is: Shortland Is (ANSP 327646); Aola, Guadalcanal (NHMUK 1952.1.4.150; ZMB); Santa Cruz Is (AM C. 29754). Occurrence in Kalimantan (Indonesian Borneo) is unconfirmed, but there are two samples in ZMB with untraceable localities in ‘East Borneo’ or ‘Southeast Borneo’. Habitat and ecology. In Bohol, Philippines, it has been reported to climb on Nypa palms in the middle and upper estuary of the Abatan River (salinity 2–10 ppt) and to be replaced by C. dohrni where salinity rises above 10 ppt (Lozouet & Plaziat 2008). It was collected by von Martens (1897 a) in slightly salty swamps with Neritina and Neritodryas, and in brackish lagoons on the trunks of the mangrove Sonneratia. Remarks. Shell morphology is highly variable and four distinctive regional forms can be distinguished. There is some apparent intergradation between forms, but their ranges are separated by either open-water gaps or regions without adequate sampling (e.g. northern New Guinea and southern Banda Sea): (1) Philippines (Fig. 5 A–K, V, W): whorls rounded; axial ribs strong and close-set; often 1–2 dark brown bands on spire contrasting with whitish ribs. This is the typical form, on which C. balteata and all its synonyms are based. (2) Sulawesi, Moluccas, Lesser Sunda Is (Fig. 5 L–P, CC, DD): whorls more flattened; axial ribs strong and closeset; often a pattern of 4–5 brown lines above periphery. (3) Western New Guinea and Aru Is (Fig. 5 Q–U): shell thin; axial ribs weak, narrow and irregular; spiral ridges present between ribs on final 2–4 whorls; often a single broad brown band above periphery. (4) Solomon Is (Fig. 5 X–BB): shell thick; axial ribs strong but sparse and remaining well developed after ventrolateral varix; usually dark brown. A single specimen from Luwuk, Sulawesi (ZMB 191.377) had a mean uncorrected pairwise distance for COI of 0.05 from specimens from the Philippines (close to the minimum between recognized species, 0.072 for the pair C. anticipata and C. reidi, Reid et al. 2013; Fig. 1), suggesting that at least forms (1) and (2) could perhaps be distinct species. The geographical range is very wide for a single species of Cerithidea and spans several range endpoints and recognized phylogeographic boundaries between sister species in the genus, which could also point to this being a species complex. On the other hand, the range extends across all the islands of eastern Indonesia and the species (or complex) has reached the Solomon Islands, unlike any other member of the genus; this could imply a wider larval dispersal, and consequently wider range, than in other congeners. Further study with denser sampling of this widespread and variable species is required. The distribution of this species covers the Philippines, eastern Indonesia, northwestern New Guinea and the Solomon Is, which are all areas of low oceanic primary productivity. Purely on the basis of this geographical pattern, this species could be described as more ‘oceanic’ than its congeners, which show typically ‘continental’ distribution patterns. This distinction has been emphasized among littorinid gastropods (e.g. Echinolittorina; Williams & Reid 2004; Reid et al. 2006) and may relate to an aspect of the ecology of planktotrophic larvae. Oceanic littorinids typically show wider distributions and extend further across the Pacific Plate than their continental congeners, and C. balteata also does so. However, C. balteata also occurs in southwestern New Guinea on the Sahul Shelf, which is an area of high oceanic productivity, so the value of the oceanic/continental distinction in this genus is uncertain. In molecular phylogenies based on the COI and 28 S rRNA genes (Reid et al. 2013; Fig. 1), this species forms a well-supported clade together with C. sinensis and C. charbonnieri. These three species should not, however, be confused: C. sinensis has a sinuous apertural margin, no peripheral keel and no ventrolateral varix; C. charbonnieri has a sinuous apertural margin, a sharp peripheral keel and a ventrolateral varix; C. balteata has a planar and expanded aperture, a moderate peripheral cord and usually a ventrolateral varix (Figs 5, 7). Their geographical distributions do not overlap. This species has been widely confused with C. tonkiniana (both being combined under the name ‘ C. ornata ’). Although their distributions do not overlap (C. tonkiniana occurring in southern Japan, China and Vietnam) and they are not phylogenetically close (Fig. 1), their shells are extremely similar. That of C. balteata is more variable, with 8–31 axial ribs on the penultimate whorl, a ventrolateral varix usually at 210–280 ° and no spiral sculpture on the upper spire whorls (Fig. 5). The shell of C. tonkiniana is slightly broader and more solid, has 11–25 axial ribs on the penultimate whorl, a varix usually at 180–220 ° and (if not eroded, which most are) shows 5 weak spiral ridges on the spire (Fig. 13 N–Y). In the western half of New Guinea, C. balteata occurs together with C. anticipata. Typical members of both species are easily separated by the lack of spiral sculpture in C. balteata. However, in the area of sympatry their discrimination can be difficult, because the local form of C. balteata does have spiral sculpture; this is usually relatively weak, the ridges are slightly irregularly spaced and do not cross the strong axial ribs (Fig. 5 Q–U). In contrast, the spiral ridges of C. anticipata are stronger, more regularly spaced and cross the weaker axial ribs to give a cancellate effect (Fig. 10 E–G). In addition, shells of C. balteata are a little narrower, more cylindrical and are more likely to show a previous apertural lip on the body whorl. Available data suggest that C. balteata occupies habitats of reduced salinity, whereas C. anticipata occurs in fully marine habitats; this distinction is likely to be maintained in regions of sympatry. This species is not collected for food in Bohol (Lozouet & Plaziat 2008)., Published as part of Reid, David G., 2014, The genus Cerithidea Swainson, 1840 (Gastropoda: Potamididae) in the Indo-West Pacific region, pp. 1-65 in Zootaxa 3775 (1) on pages 18-23, DOI: 10.11646/zootaxa.3775.1.1, http://zenodo.org/record/285731, {"references":["Adams, A. (1855) A monograph of Cerithidea, a genus of Mollusca, with descriptions of several new species, from the collection of High Cuming, Esq,: to which are added, descriptions of two new species of Colina, and one of Donax. Proceedings of the Zoological Society of London, 1854, 83 - 87. http: // dx. doi. org / 10.1111 / j. 1469 - 7998.1854. tb 07236. x","Hidalgo, J. G. (1904 - 1905) Catalogo de los moluscos testaceos de las Islas Filipinas, Jolo y Marianas. I. - Moluscos marinos. Gaceta de Madrid, Madrid, xvi + 408 pp.","Reid, D. G., Claremont, M., Smith, L., Shamoto, M., Glaubrecht, M. & Ozawa, T. (2013) Mosaics in the mangroves: allopatric diversification of tree-climbing mudwhelks (Gastropoda: Potamididae: Cerithidea) in the Indo-West Pacific. Biological Journal of the Linnean Society, 110, 564 - 580.","Cecalupo, A. (2005) Elenco della famiglia Cerithiidae Ferussac, 1822 (Prosobranchia). Nomenclatura delle specie conosciute o poco note e relativa revisione sistematica. Fossili e Attuali. Vol. I. Quaderni della Civica Stazione Idrobiologica di Milano, 26, 1 - 366.","Casto de Elera, R. P. (1896) Catalogo Sistematico de Toda la Fauna de Filipinas, Vol. 3. Moluscos y Radiados. Colegio de Santo Tomas, Manila, 942 + lxiv pp.","Sowerby, G. B. II (1866) Monograph of the genus Cerithidea. In: Reeve, L. A. (Ed.), Conchologia Iconica. Vol. 15. L. A. Reeve, London, pls 1 - 4.","Sowerby, G. B. II (1855) Monograph of the genus Cerithium, Adanson. In: Sowerby, G. B. II. (Ed.), Thesaurus Conchyliorum, or Figures and Descriptions of Recent Shells. Vol. 2. part 16. Sowerby, London, pp. 847 - 899.","Tryon, G. W. (1887) Manual of Conchology. Vol. 9. Tryon, Philadelphia, 488 pp.","Martens, E. von (1897 a) Suss- und Brackwasser-Mollusken des Indischen Archipels. In: Weber, M. (Ed.), Zoologische Ergebnisse einer Reise in Niederlandisch Ost-Indien, Vol. 4. E. J. Brill, Leiden, pp. 1 - 331.","Kobelt, W. (1890 b) Eine neue Cerithidea. Nachrichtsblatt der deutschen Malakozoologischen Gesellschaft, 1890, 60 - 61.","Schepman, M. M. (1909) The Prosobranchia of the Siboga Expedition. Part 2. Taenioglossa and Ptenoglossa. Siboga-Expeditie, 49, 109 - 231.","Oostingh, C. H. (1925) Report on a collection of Recent shells from Obi and Halmahera (Moluccas). Communications from the Geological Institute of the Agricultural University, Wageningen (Holland), 9, 1 - 362.","Lozouet, P. & Plaziat, J. - C. (2008) Mangrove Environments and Molluscs. Abatan River, Bohol and Panglao Islands, Central Philippines. ConchBooks, Hackenheim, 160 pp.","Reid, D. G., Dyal, P., Lozouet, P., Glaubrecht, M. & Williams, S. T. (2008) Mudwhelks and mangroves: the evolutionary history of an ecological association (Gastropoda: Potamididae). Molecular Phylogenetics and Evolution, 47, 680 - 699. http: // dx. doi. org / 10.1016 / j. ympev. 2008.01.003","Van Benthem Jutting, W. S. S. (1956) Systematic studies on the non-marine Mollusca of the Indo-Australian archipelago. V. Critical revision of the Javanese freshwater gastropods. Treubia, 23, 259 - 477.","Say, T. (1825) Descriptions of some new species of fresh water and land snails of the United States. Journal of the Academy of Natural Sciences of Philadelphia, 5, 119 - 131.","Cecalupo, A. (2006) Elenco della famiglia Cerithiidae Ferussac, 1822 (Prosobranchia). Nomenclatura delle specie conosciute o poco note e relativa revisione sistematica. Attuali. Vol. II. Quaderni della Civica Stazione Idrobiologica di Milano, 27, 1 - 373.","Iredale, T. (1929) Queensland molluscan notes, no. 1. Memoirs of the Queensland Museum, 9, 261 - 297.","Petit, R. E. (2009) George Brettingham Sowerby, I, II & III: their conchological publications and molluscan taxa. Zootaxa, 2189, 1 - 218.","Philippi, R. A. (1848) testaceorum novorum centuria (continuatio). Zeitschrift fur Malakozoologie, 1848, 17 - 27.","Williams, S. T. & Reid, D. G. (2004) Speciation and diversity on tropical rocky shores: a global phylogeny of snails of the genus Echinolittorina. Evolution, 58, 2227 - 2251. http: // dx. doi. org / 10.1554 / 03 - 565","Reid, D. G., Lal, K., Mackenzie-Dodds, J., Kaligis, F., Littlewood, D. T. J. & Williams, S. T. (2006) Comparative phylogeography and species boundaries in Echinolittorina snails in the central Indo-West Pacific. Journal of Biogeography, 33, 990 - 1006. http: // dx. doi. org / 10.1111 / j. 1365 - 2699.2006.01469. x"]}

Cerithidea andamanensis new species (Figures 11, 12 K–P) Potamides (Cerithidea) quadratus —von Martens, 1887: 169 (not Sowerby, 1866 = C. quoyii). Cerithidea n. sp. B Reid et al., 2013: fig. 2 (map). Types. Holotype NHMUK 1887.3. 10.24 (Fig. 12 K–N) and 4 dry paratypes NHMUK 1887.3.10.25– 28, King I. [Kadan Kyun], Mergui [Myeik] Archipelago, Burma. Etymology: from the known distribution in the Andaman Sea. Taxonomic history. This species has only recently been recognized, its existence having been noted in the biogeographic study of Reid et al. (2013). The identification by von Martens’ (1887) is based on the same material from Mergui (collected by J. Anderson) which now forms the type collection. Diagnosis. Shell: spire straight, whorls rounded, periphery weakly angled; aperture flared, anterior canal and apertural projection well developed; 16–22 axial ribs on penultimate whorl, 7–10 after ventrolateral varix; ventrolateral varix an enlarged rib at 240–250 °; 5–6 spiral ridges on spire, 4–5 ridges above periphery on last whorl, spiral ridges form nodules where they cross axial ribs but are weak in between; brown with darker spiral cords. Andaman Sea. Material examined. 11 lots. Shell (Fig. 12 K–P): H = 26–39.2 mm. Shape elongated conical (H/B = 2.29–2.47, SH = 3.00– 3.43); decollate, 9 whorls remaining; spire whorls rounded, suture distinct; spire profile straight; periphery weakly angled; thickness moderate. Adult lip flared, thickened; apertural margin planar in side view; strong anterior projection adjacent to deep notch of anterior canal. Sculpture on spire of straight to slightly curved (opisthocyrt) rounded axial ribs, interspaces 1–2 times width of ribs, 16–22 ribs on penultimate whorl, 7–10 ribs after ventrolateral varix (i.e. only slightly more distant than on spire), remaining strong throughout final whorl, only faint axial wrinkles on base; spire whorls with 5–6 spiral ridges, forming small nodules where they cross axial ribs but weakly developed in spaces between axial ribs, spiral interspaces 1–1.5 times width of ridges, no interpolated threads, 4–5 spiral ridges above periphery on last whorl; overall effect is of nodulose axial ribs; base with 12–14 spiral ridges, outermost is peripheral cord of same size as primary ridges above. Ventrolateral varix a strongly enlarged rib at 240–250 °, forming an anteriorly projecting boss at periphery. Surface with fine spiral microstriae on periostracum, strongest on spiral ridges. Colour: brown, spiral ridges darker brown, shoulder ridge sometimes pale; aperture pale brown, spiral lines showing through. Animal: unknown. Range (Fig. 11): W Thailand, Andaman Is, NW Malaysia, NE Sumatra. Records: India: Andaman Is (NHMUK 1907.8.30.267). Burma: King I. [Kadan Kyun], Mergui [Myeik] Arch. (NHMUK 1887.3.10.24; ZMB). Thailand: Ranong (USNM 777649); Ao Nam-Bor, Phuket I. (NHMUK 20130250); Kantang, Trang Prov. (RMNH). Malaysia: Langkawi, Kedah (T. Goulding & B. Dayrat). Indonesia: Pantai Cermin, Deli, Sumatra (RMNH 175798). Habitat and ecology. At Phuket I., Thailand, a specimen was collected in the landward fringe of a marine mangrove forest. Remarks. No molecular data are available. However, the shell characters of the available specimens differ in a consistent way from the two other members of the C. quoyii group and these differences are even more pronounced than those that distinguish the other two from each other (Table 1). The distributional range of the new species approaches closely to that of C. quoyii in the Strait of Malacca (Fig. 11). In western Malaysia the southernmost record of the new species is from Langkawi, while C. quoyii has been recorded from Kelang, 420 km to the south. One sample from northeastern Sumatra (RMNH 175798) contains four typical C. quoyii and one of the form described here, indicating that they remain distinct where their ranges meet. It is therefore predicted that these shells represent a genetically distinct species. The most significant characters are the rounded whorls, 5–6 weak spiral ridges on the spire whorls and the strong, numerous axial ribs continuing after the ventrolateral varix. The Andaman Sea is an area of lower nutrient status and clear water, implying that this species is more ‘oceanic’ in its habitat range than C. quoyii, which is found in turbid conditions on the nutrient-rich continental shores of Southeast Asia. The shell is similar to that of some forms of the variable C. anticipata from Australia (Fig. 10 A–P), but the periphery is more angular and the peristome consequently slightly quadrate, rather than round. The ranges of these two taxa are widely disjunct., Published as part of Reid, David G., 2014, The genus Cerithidea Swainson, 1840 (Gastropoda: Potamididae) in the Indo-West Pacific region, pp. 1-65 in Zootaxa 3775 (1) on page 42, DOI: 10.11646/zootaxa.3775.1.1, http://zenodo.org/record/285731, {"references":["Martens, E. von (1887) List of the shells of Mergui and its archipelago, collected for the trustees of the Indian Museum, Calcutta, by Dr. John Anderson, F. R. S., Superintendent of the Museum. Journal of the Linnean Society of London, Zoology, 21, 155 - 219. http: // dx. doi. org / 10.1111 / j. 1096 - 3642.1887. tb 00974. x","Sowerby, G. B. II (1866) Monograph of the genus Cerithidea. In: Reeve, L. A. (Ed.), Conchologia Iconica. Vol. 15. L. A. Reeve, London, pls 1 - 4.","Reid, D. G., Claremont, M., Smith, L., Shamoto, M., Glaubrecht, M. & Ozawa, T. (2013) Mosaics in the mangroves: allopatric diversification of tree-climbing mudwhelks (Gastropoda: Potamididae: Cerithidea) in the Indo-West Pacific. Biological Journal of the Linnean Society, 110, 564 - 580."]}

Genus Cerithidea Swainson, 1840 Cerithidea Swainson, 1840: 198, 203, 342 (type by subsequent designation (Makiyama 1936) Melania lineolata Gray in Griffith & Pidgeon, 1833 = C. obtusa). Phaenommia Mörch, 1860: 80 (type by monotypy Cerithidea charbonieri (sic) = Cerithium charbonnieri Petit, 1851). Potamides (Aphanistylus) Fischer, 1884: 682 (type by monotypy Cerithidea charbonnieri = Cerithium charbonnieri Petit, 1851). Taxonomic history. When introducing the new genus Cerithidea, Swainson (1840) gave only two included species, Melania lineolata (= C. obtusa, herein) and the fossil Cerithium fragilis Brogniart. The subsequent history of the designation of the former as the type of the genus has been discussed by Bequaert (1942 b), who concluded that the first valid designation (of C. obtusa) was that of Pilsbry & Harbison (1933). This conclusion was accepted by Houbrick (1984) but, as pointed out by Petit & Coan (2008), this designation was invalid, because it did not mention either of the originally included taxa, so that the first valid designation was that of Makiyama (1936). Two generic names have been based on C. charbonnieri, by Mörch (1860) and Fischer (1884). Both authors cited the long optic peduncles and short or invisible tentacles in the figure by H. Adams & A. Adams (1854: pl. 31, fig. 2). In fact the tentacles of living and preserved Cerithidea species are often folded back and not immediately visible (von Martens 1897 a: pl. 10, fig. 5), and the peduncles are not unusually long in this species. The shell of C. charbonnieri is unusually smooth for the genus, with a strong peripheral carina. Aphanistylus has occasionally been used as a subgenus (Fischer 1884; Kobelt 1890 a; Casto de Elera 1896; Dautzenberg 1899) and rarely as a genus (Wattebled 1886; Morlet 1889), for one or more of C. weyersi, C. charbonnieri, C. sinensis and C. balteata. Thiele (1929) remarked that the subgenus Aphanistylus “might be hardly different” from Cerithidea s. s., but nevertheless later replaced it with the earlier name Phaenommia (Thiele 1931). Wenz (1939) formally listed Aphanistylus and Phaenommia in the synonymy of Cerithidea. The three species C. charbonnieri, C. sinensis and C. balteata do form a well-supported molecular clade (Fig. 1). Nevertheless, there is no known morphological synapomorphy that could justify the continued use of this subgeneric unit. The generic name Cerithidea is here used only for those potamidids from the Indo-West Pacific that have a close, or obligate, association with mangrove trees and halophytic vegetation. This concept of Cerithidea is that of Reid et al. (2008) and before this the generic classification of its species had a complex history. Cerithidea was established by Swainson (1840) and included as a genus in the classification by Gray (1847). This early concept of the genus included American species (assigned to Cerithideopsis by Reid et al. 2008) as well as the Indo-West Pacific members (H. Adams & A. Adams 1854; Adams 1855; Sowerby 1866). The smaller, mud-dwelling Indo- West Pacific potamidids (assigned to Cerithideopsilla by Reid et al. 2008) were classified in Tympanotonos by these early authors, while Pirenella contained the diverse forms of the polymorphic P. conica from the Mediterranean and Indian Ocean. Some nineteenth-century authors maintained a much broader generic concept and included all these potamidids in Cerithium (Kiener 1841 –1842; Sowerby 1855; Reeve 1860), sometimes with Cerithidea, Tympanotonos and Pirenella as subgenera (Chenu 1859; Kobelt 1890 –1895). Fischer (1884) established the use of Potamides (of which the type was an Oligocene fossil) with these three potamidid groups as subgenera, and this scheme was widely followed (Tryon 1887; von Martens 1897 a; Cossmann 1906, but treating the Recent Tympanotonos s. s. as a separate genus). The prevailing twentieth-century concept of Cerithidea dates from Thiele (1929), who introduced Cerithideopsis as a subgenus, with sections Cerithideopsis s. s. for American species and Cerithideopsilla for Indo- West Pacific ones, while continuing to recognize Pirenella as a genus. This scheme was modified only slightly by Wenz (1938), who established the tripartite division of Cerithidea in three subgenera: Cerithidea s. s., Cerithideopsis and Cerithideopsilla. This was followed by most subsequent authors including Van Regteren Altena (1940), Bequaert (1942 a) and Brandt (1974), and was supported by a detailed anatomical study by Houbrick (1984). Pirenella continued to be treated as a genus, but was synonymized with Potamides by Lozouet (1986). As a result of molecular phylogenetic analysis and a review of the shell characters of living and fossil genera, Reid et al. (2008) proposed a new classification of the Potamididae. The three former subgenera Cerithidea, Cerithideopsis and Cerithideopsilla were raised to generic rank, the enigmatic C. largillierti from the western Pacific was classified in Cerithideopsis for the first time and ‘ Potamides ’ conicus was shown to be a member of Cerithideopsilla. Diagnosis. Shell: large (to 61 mm), delicate to solid, elongate, spire decollate; sculpture of axial ribs, one developed as prominent ventrolateral varix on body whorl (rarely weak or absent) and weakening thereafter; varices almost always absent on spire whorls; spiral ridges present on base, present or absent above periphery; aperture circular, peristome flared and thickened in adult, planar or slightly sinuous, with short anterior canal and slight posterior groove; columella straight or slightly twisted, lacking folds; colour often with spiral bands. Radula: rachidian tooth narrow with small central cusp; outer marginal lacks cusps and has wide flange on outer side of basal shaft (Houbrick 1984: 16). Remarks. The diagnostic characters of the shells of the three similar potamidid genera Cerithidea, Cerithideopsis and Cerithideopsilla were summarized by Reid et al. (2008). In Cerithideopsis there are 2–3 spiral cords on the earliest spire whorls, which usually become obsolete or may increase to 5; there are numerous axial ribs that terminate at the strong spiral cord demarcating the base; varices are scattered over the spire and the ventrolateral varix is absent. In Cerithideopsilla there are 3 spiral cords, numerous axial ribs and no varices on the spire whorls; the ventrolateral varix is usually present and the anterior canal of the aperture is well defined by a basal projection of the aperture. The spire is not decollate in either of these two genera, as it is in Cerithidea. The shell of Cerithidea is typically of light construction, as expected in relation to the habit of climbing onto trunks and vegetation, where the animals rest, attached only by mucus at the apertural lip, between periods of feeding activity on the substrate at low tide. One of the largest is C. obtusa, which is also more thick-shelled than other species; it rests on tree bases and even on the ground, where its relatively solid shell may be resistant to attack by predators such as large crabs and molluscivorous fish. Large samples of several species show dimorphism in shell size, which is possibly related to sex (e.g. Fig. 8 G, H), although this remains to be established. All species are decollate, the apical 5–10 mm of the spire being lost (e.g. Figs 3 K, L, 15 O, P). This is a gradual process of dissolution of the delicate early whorls, which appears to continue throughout life, and the resulting opening is sealed by successive convex plugs of shell material secreted at the tip of the visceral mass. Dissolution does not appear to be an endogenous process connected with conservation of calcium carbonate or neutralization of tissue acidity during anaerobic respiration (cf. Vermeij 1973). Instead, it is likely to be the result of erosion by acidic conditions in the detritus-rich sediments on which the snails crawl to feed at low tide. The ventral surface of the adult shell, in prolonged contact with the sediment, sometimes shows signs of dissolution that may even remove the surface sculpture (Figs 3 N–P, 12 H). Similarly, dissolution pits may be seen wherever the periostracum is damaged. Decollation may, nevertheless, be adaptive. It is a derived character not seen to such an extreme in other potamidids (Reid et al. 2008) and might reduce the weight of the shell, or reduce vulnerability to predatory attack by removing the more easily damaged apical whorls. Growth is determinate and the final whorl is modified by reduction of axial sculpture, the formation of a single strong axial rib (the ‘ventrolateral varix’) and a final flaring and thickening of the aperture (compare juvenile and adult in Figs 12 O, K, 14 I, A). This growth pattern is typical of cerithioideans and has been considered adaptive in relation to stability on soft substrates and protection against predatory crabs that attack the aperture and ‘peel’ the shell (Sälgeback & Savazzi 2006). Vermeij (1973) pointed out that in tree-climbing potamidids the form of the shell is a compromise between achieving stability on sediment and on a hard substrate. The flared aperture generally has a planar margin and is inclined at an angle to the axis of coiling (i.e. opisthocline) so that the shell opening can be held closely against a flat surface such as the trunk of a tree (Fig. 12 T, V); in the potamidid Terebralia sulcata this has been considered as an adaptation against desiccation (Sälgeback & Savazzi 2006), but in Cerithidea it is likely that this shape also functions to improve attachment by dried mucus. In five species (C. sinensis, C. charbonnieri, C. tonkiniana, C. rhizophorarum, C. moerchii) the apertural margin is slightly sinuous in side view so that the aperture cannot be applied tightly to a flat surface (Fig. 7 G, Q). Three of these five species (C. sinensis, C. tonkiniana, C. moerchii) extend to the highest latitudes in the entire genus, beyond the mainly tropical range of mangrove trees, and can be found attached to salt marsh plants and reeds, so this form of the aperture could be functional in relation to attachment to stems of narrow diameter. Whether the ventrolateral varix improves stability by preventing rolling is doubtful, because its position is rather variable (e.g. Fig. 12 Q–S, W–Y, AA–CC); stability would only be enhanced if this varix were always close to 270 ° behind the apertural edge. Instead, it is likely that the varix serves mainly as a second line of defence against predators that break the thickened aperture and peel away the thin body whorl. Evidence of such damage that stops at the ventrolateral varix is frequent in Cerithidea but, as observed in Terebralia (Sälgeback & Savazzi 2006), sublethal breakages do not extend past this barrier, presumably because they are always lethal. In Cerithidea the ventrolateral varix is not only thickened, but also bears two internal teeth (pers. obs. in C. moerchii). In some specimens this varix appears to be formed by the fusion of two axial ribs. The anatomy and radula of Cerithidea have been described by Houbrick (1984, 1986). A notable peculiarity is the presence of a pallial eye on the mantle margin, which protrudes into the anterior canal of the shell when the animal is at rest. In C. obtusa the eye has an epithelial lens and cup-shaped cornea (Pflugfelder 1930) and pallial eyes of varying complexity are found in Cerithideopsis, Cerithideopsilla and other cerithioideans (Houbrick 1984). As expected in snails that live mainly out of water, the gill and osphradium are reduced and the mantle cavity is vascularized (Houlihan 1979; Houbrick 1984). The living animal is sometimes brightly coloured, as first recorded by Adams (in Adams & Reeve 1848; see also Fukuda 2000; Fig. 2). In Southeast Asia several common species (especially C. obtusa and C. quoyii) are of local economic importance and sold in food markets (Brandt 1974; Poutiers 1998; Hamli et al. 2013). In the following accounts of the species, the arrangement follows an approximate phylogenetic sequence. The COI phylogeny (Reid et al. 2013; Fig. 1) identifies the following monophyletic groups: (1) C. weyersi; (2) C. balteata, C. sinensis, C. charbonnieri; (3) C. obtusa; (4) C. anticipata, C. reidi, C. quoyii, C. dohrni; (5) C. decollata; (6) C. tonkiniana, C. moerchii. The species C. houbricki, C. andamanensis and C. rhizophorarum are known only from shell material and have been added to groups 1, 4 and 6 respectively, on the basis of shell resemblance., Published as part of Reid, David G., 2014, The genus Cerithidea Swainson, 1840 (Gastropoda: Potamididae) in the Indo-West Pacific region, pp. 1-65 in Zootaxa 3775 (1) on pages 9-13, DOI: 10.11646/zootaxa.3775.1.1, http://zenodo.org/record/285731, {"references":["Swainson, W. (1840) A Treatise on Malacology. Longman, Orme, Brown, Green & Longmans, London, viii + 419 pp.","Makiyama, J. (1936) The Meisen Miocene of North Korea. Memoirs of the College of Science, Kyoto Imperial University, Series B, 11, 193 - 228.","Griffith, E. & Pidgeon, E. (1833 - 1834) The Mollusca and Radiata. In: Griffith, E., (Ed.) The Animal Kingdom arranged in conformity with its organization, by the Baron Cuvier, Vol. 12. Whittaker & Co., London, viii + 601 pp.","Morch, O. A. L. (1860) Beitrage zur Molluskenfauna Central-Amerika's. Malakozoologische Blatter, 7, 66 - 106.","Petit de la Saussaye, M. S. (1851) Description de coquilles nouvelles. Journal de Conchyliologie, 2, 259 - 269.","Bequaert, J. C. (1942 b) Random notes on American Potamididae. Nautilus, 56, 20 - 30.","Pilsbry, H. A. & Harbison, A. (1933) Notes on the Miocene of southern New Jersey. Proceedings of the Academy of Natural Sciences of Philadelphia, 85, 107 - 120.","Houbrick, R. S. (1984) Revision of higher taxa in genus Cerithidea (Mesogastropoda: Potamididae) based on comparative morphology and biological data. American Malacological Bulletin, 2, 1 - 20.","Petit, R. E. & Coan, E. V. (2008) The molluscan taxa made available in the Griffith & Pidgeon (1833 - 1834) edition of Cuvier, with notes on the editions of Cuvier and on Wood's Index Testaceologicus. Malacologia, 50, 219 - 264. http: // dx. doi. org / 10.4002 / 0076 - 2997 - 50.1.219","Martens, E. von (1897 a) Suss- und Brackwasser-Mollusken des Indischen Archipels. In: Weber, M. (Ed.), Zoologische Ergebnisse einer Reise in Niederlandisch Ost-Indien, Vol. 4. E. J. Brill, Leiden, pp. 1 - 331.","Kobelt, W. (1890 b) Eine neue Cerithidea. Nachrichtsblatt der deutschen Malakozoologischen Gesellschaft, 1890, 60 - 61.","Casto de Elera, R. P. (1896) Catalogo Sistematico de Toda la Fauna de Filipinas, Vol. 3. Moluscos y Radiados. Colegio de Santo Tomas, Manila, 942 + lxiv pp.","Dautzenberg, P. (1899) Contribution a la faune malacologique de Sumatra (recoltes de M. J. - L. Weyers). Annales de la Societe Royale Malacologique de Belgique, 34, 1 - 26.","Wattebled, G. (1886) Description de mollusques inedits de l'Annam. Recolte du capitaine Dorr aux environs de Hue. Journal de Conchyliologie, 34, 54 - 71.","Morlet, L. (1889) Catalogue des coquilles recueillis, par M. Pavie, dans le Cambodge et le Royaume de Siam, et description d'especes nouvelles. Journal de Conchyliologie, 37, 121 - 199.","Thiele, J. (1929, 1931) Handbuch der Systematischen Weichtierkunde. Parts 1, 2. Gustav Fischer, Jena, 777 pp.","Reid, D. G., Dyal, P., Lozouet, P., Glaubrecht, M. & Williams, S. T. (2008) Mudwhelks and mangroves: the evolutionary history of an ecological association (Gastropoda: Potamididae). Molecular Phylogenetics and Evolution, 47, 680 - 699. http: // dx. doi. org / 10.1016 / j. ympev. 2008.01.003","Gray, J. E. (1847) A list of the genera of recent Mollusca, their synonyma and types. Proceedings of the Zoological Society of London, 1847, 129 - 219.","Adams, A. (1855) A monograph of Cerithidea, a genus of Mollusca, with descriptions of several new species, from the collection of High Cuming, Esq,: to which are added, descriptions of two new species of Colina, and one of Donax. Proceedings of the Zoological Society of London, 1854, 83 - 87. http: // dx. doi. org / 10.1111 / j. 1469 - 7998.1854. tb 07236. x","Sowerby, G. B. II (1866) Monograph of the genus Cerithidea. In: Reeve, L. A. (Ed.), Conchologia Iconica. Vol. 15. L. A. Reeve, London, pls 1 - 4.","Kiener, L. - C. (1841 - 1842) Genre Cerite. (Cerithium, Adanson.) In: Kiener, L. - C. (Ed.), Species General et Iconographie des Coquilles Vivantes. Rousseau & Bailliere, Paris, 104 pp.","Sowerby, G. B. II (1855) Monograph of the genus Cerithium, Adanson. In: Sowerby, G. B. II. (Ed.), Thesaurus Conchyliorum, or Figures and Descriptions of Recent Shells. Vol. 2. part 16. Sowerby, London, pp. 847 - 899.","Reeve, L. A. (1860) Elements of Conchology. Vol. 1. Reeve, London, 260 pp.","Chenu, J. C. (1859) Manuel de Conchyliologie. Vol. 1. G. Masson, Paris, vii + 508 pp.","Tryon, G. W. (1887) Manual of Conchology. Vol. 9. Tryon, Philadelphia, 488 pp.","Cossmann, M. (1906) Essais de Paleoconchologie Comparee, Vol. 7. Rudeval, Paris, 261 pp.","Wenz, W. (1938) Gastropoda. Teil I: Allgemeiner Teil und Prosobranchia. In: Schindewolf, O. H. (Ed.), Handbuch der Palaozoologie. Vol. 6. Borntraeger, Berlin, xii + 948 pp.","Van Regteren Altena, C. O. (1940) A revision of Cerithidea (Cerithideopsilla) cingulata (Gmelin) and some related species (Mollusca, Gastropoda). Zoologische Mededelingen, 22, 211 - 222.","Bequaert, J. C. (1942 a) Cerithidea and Batillaria in the western Atlantic. Johnsonia, 5, 1 - 11.","Brandt, R. A. M. (1974) The non-marine aquatic Mollusca of Thailand. Archiv fur Molluskenkunde, 105, 1 - 423.","Lozouet, P. (1986) Redefinition des genres Potamides et Pirenella (Gastropoda, Prosobranchia) a partir des especes actuelles et fossiles: implications phyletiques et biogeographiques. Annales de Paleontologie, 72, 163 - 210.","Vermeij, G. J. (1973) Molluscs in mangrove swamps: physiognomy, diversity, and regional differences. Systematic Zoology, 22, 609 - 624. http: // dx. doi. org / 10.2307 / 2412965","Salgeback, J. & Savazzi, E. (2006) Constructional morphology of cerithiform gastropods. Paleontological Research, 10, 233 - 259. http: // dx. doi. org / 10.2517 / prpsj. 10.233","Houbrick, R. S. (1986) Cerithidea reidi, spec. nov., from Western Australia. Veliger, 28, 280 - 286.","Pflugfelder, O. (1930) Das Mantelauge von Potamides obtusus Lam. Zoologischer Anzeiger, 89, 276 - 283.","Houlihan, D. F. (1979) Respiration in air and water of three mangrove snails. Journal of Experimental Marine Biology and Ecology, 41 (2), 143 - 161. http: // dx. doi. org / 10.1016 / 0022 - 0981 (79) 90049 - 2","Adams, A. & Reeve, L. (1848 - 1850) Mollusca. In: Adams, A. (Ed.), The Zoology of the Voyage of H. M. S. Samarang. Reeve, Benham & Reeve, London, x + 87 pp.","Fukuda, H. (2000) Gastropods I: general view. In: Sato, M. (Ed.), Life in Ariake Sea: Biodiversity in Tidal Flats and Estuaries. Kaiyu-sha, Tokyo, pp. 213 - 252.","Poutiers, J. M. (1998) Gastropods. In: Carpenter, K. E. & Niem, V. H. (Eds.), The Living Marine Resources of the Western Central Pacific. Vol. 1. FAO, Rome, pp. 363 - 648.","Hamli, H., Idris, M. H., Abu Hena, M. K., Wong, S. K. & Arshad, A. (2013) Checklist and habitat of edible gastropods from Sarawak, Malaysia. Journal of Fisheries and Aquatic Sciences, 8, 412 - 418. http: // dx. doi. org / 10.3923 / jfas. 2013.412.418","Reid, D. G., Claremont, M., Smith, L., Shamoto, M., Glaubrecht, M. & Ozawa, T. (2013) Mosaics in the mangroves: allopatric diversification of tree-climbing mudwhelks (Gastropoda: Potamididae: Cerithidea) in the Indo-West Pacific. Biological Journal of the Linnean Society, 110, 564 - 580."]}

Cerithidea quoyii (Hombron & Jacquinot, 1848) (Figures 2 D, 11, 12B–J) Cerithium obtusum — Adams & Reeve, 1848: 43 –44, pl. 13, fig. 3 a, b (not Lamarck, 1822). Cerithium quoyii Hombron & Jacquinot, 1848: pl. 23, figs 6, 7 (Singapour [Singapore]; lectotype here designated Hombron & Jacquinot 1848: pl. 23, fig. 6). Rousseau, 1854: 97. Cerithidea quoyii — Reid et al., 2013: figs 1 (phylogeny), 2 (map). Cerithium truncatum — Reeve, 1860: 124 (not Gray in Griffith & Pidgeon, 1833; not Griffith & Pidgeon, 1834; description is a copy of Adams & Reeve, 1848: 43 –44). Cerithidea quadrata G.B. Sowerby II, 1866: Cerithidea sp. 5, pl. 1, fig. 5 (Malacca [Melaka, Malaysia]; 3 syntypes NHMUK 20130228, Fig. 12 D–G, seen). Tapparone-Canefri, 1874: 144 –145. Morlet, 1889: 144. Dautzenberg & Fischer, 1906: 410 – 411 (in part, includes C. dohrni). Van Benthem Jutting, 1956: 435 –436, fig. 107. Poutiers, 1998: 454, fig. (in part, includes C. dohrni, C. andamanensis). Swennen et al., 2001: 111, fig. 294. Dharma, 2005: 92, pl. 21, fig. 4 a, b. Ng et al., 2008: 72 (living animal). Reid et al., 2008: 680 –699, figs 1, 2 (phylogeny). Mujiono, 2009: 54, fig. 1 G. Hamli et al., 2013: 412 –418, fig. 2 d. Potamides (Cerithidea) obtusa var. quadrata — Tryon, 1887: 161, pl. 33, fig. 63. Cerithium (Cerithidea) quadratum — Kobelt, 1890 a: 45 –46, pl. 9, fig. 8 (Cecalupo 2005: pl. 31, fig. 7). Potamides (Cerithidea) quadratus —von Martens, 1897 a: 187 –188, pl. 9, fig. 23; pl. 10, fig. 4 (head) (in part, includes C. anticipata). Potamides quadratum — Prashad, 1921: 495 –496. Cerithidea (Cerithidea) quadrata — Brandt, 1974: 193, pl. 14, fig. 53. Thach, 2007: 60, pl. 9, fig. 182. Potamides (Cerithidea) rhizophorarum — Tryon, 1887: 162, pl. 33, fig. 62 (in part, includes C. rhizophorarum; not A. Adams, 1855; as rhizoporarum). Cerithidea rhizophorarum — Van Regteren Altena, 1945: 145 (synonymy fide Van Benthem Jutting 1956; not A. Adams, 1855) Cerithidea obtusa —Ng & Sivasothi, 1999: 106, figs (not Lamarck, 1822). Cerithidea (Cerithideopsis) pliculosa — Cecalupo, 2005: 316, pl. 31, fig. 7 (not Menke, 1829). Cerithidea (Aphanistylus) charbonnieri — Cecalupo, 2006: 133, 186, 233 (not Petit de la Saussaye, 1851; in part, includes C. charbonnieri). Taxonomic history. It is curious that the valid name for this species, introduced by Hombron & Jacquinot in 1848 (see Taxonomic History of C. anticipata for discussion of date of publication) has been so long neglected. It was erroneously listed in the synonymy of C. rhizophorarum by Tryon (1887). It was mentioned under C. quadrata by von Martens (1897 a), perhaps unaware of its priority; it does not appear to be a preoccupied name. Since then it has not been mentioned until resurrected by Reid et al. (2013). Despite this relative obscurity, the conditions for reversal of precedence (ICZN 1999: Art. 23.9) in favour of the more widely-used C. quadrata are not met in this case. There is a specimen labelled as a syntype of Cerithium quoyii in MNHN (MNHN 25693; Fig. 12 A), but this is not the shell figured by Hombron & Jacquinot (1848: pl. 23, figs 6, 7) and it bears a label ‘Borneo’, whereas the type locality was given as ‘Singapour’. Furthermore, the shell more closely resembles C. dohrni from the Philippines, so its identification as a syntype is not accepted. The original figure and type locality are, however, sufficient to define the species unambiguously. For most of its history this species has been known under the name C. quadrata. It is not clear why Sowerby (1866) ignored the earlier name, when he considered another of Hombron & Jacquinot’s (1848) new taxa, C. kieneri, to be valid. Diagnosis. Shell: broad, spire convex, whorls flattened, periphery angled; aperture flared, anterior canal and apertural projection well developed; 14–32 axial ribs on penultimate whorl, 3–6 weakening ribs after ventrolateral varix; ventrolateral varix an enlarged rib at 240–270 °; 7 spiral cords on spire, 6–9 cords and threads above periphery on last whorl; brown with darker spiral cords. Southeast Asia, Borneo, Java. COI GenBank HE 680230, HE 680231, HE 680233 –680235, AM 932771, AM 932772. Material examined. 58 lots. Shell (Fig. 12 A–J): H = 24.1–45 mm (52 mm, Brandt 1974). Shape elongated conical, usually relatively broad (H/B = 2.11–2.69, SH = 2.37–3.17); decollate, 6–9 whorls remaining; spire whorls flattened to weakly rounded, suture indistinct; spire profile usually convex; periphery usually strongly angled; thin to moderate thickness. Adult lip flared, slightly thickened; apertural margin planar in side view; strong anterior projection adjacent to deep notch of anterior canal. Sculpture on spire of straight to slightly curved (opisthocyrt) axial ribs, narrowly rounded, interspaces 1–1.5 times width of ribs, rarely 1 rib enlarged as a varix on spire, 14–32 ribs on penultimate whorl, 3– 6 distant ribs after ventrolateral varix, weak or absent on final 0.25 whorl, only faint axial wrinkles on base; spire whorls with 7 spiral cords, approximately equal in width to interspaces, often increased by interpolation of 1–3 narrow threads on last 1–2 whorls to give 6–9 spiral cords and threads above periphery on last whorl; base with (10) 12–15 spiral threads, outermost is peripheral cord of same size as primary cords above and raised as a keel. Ventrolateral varix a strongly enlarged rib at 240–270 °, forming an anteriorly projecting boss at periphery. Surface with fine spiral microstriae on periostracum, strongest on spiral cords. Colour: brown, spiral cords darker brown except for 2–3 pale cords at shoulder (giving appearance of pale spiral band); aperture pale brown, spiral lines showing through. Animal (Fig. 2 D): Head and base of tentacles pinkish grey with cream spots; anterior half of snout blackish, sometimes with a few yellow spots, if black pigment is less intense then snout and tentacle bases appear more strongly pink; tentacles pale grey with black rings; sides of foot grey, blackish anteriorly, with small yellowish cream spots; sole of foot grey, pinkish towards margin; mantle pale pinkish grey (based on ethanol-preserved specimens). Adams & Reeve (1848) described and illustrated a light brown snout with three longitudinal, broad, opaque yellow lines, the central one longest and reaching nearly to the tip where it is bifid, the foot light pinkish brown mottled with dark brown and the sole lilac. Range (Fig. 11): S Vietnam, E Thailand, Malaysia, Borneo, Java, S Sulawesi. Records: Vietnam: Ninh Phu, 25 km N Nha Trang (NHMUK 20010373); Vung Tau (ANSP 330752). Cambodia: Kampot (Morlet 1889). Thailand: Chantaburi R. (RMNH; AM C. 79266; USNM 794072); Koh Taluei (USNM 361191). Malaysia: Leban Condong, Pahang (AM C. 124469); Klang, Selangor (NHMUK); Koyong Hujan, Sarawak (USNM 673304); Uglam Hujung, Kudat, Sabah (NHMUK 20130247); Sandakan, Sabah (USNM 666744; ANSP 295506). Brunei: Sungai Ayam Ayam, Brunei Bay (NHMUK). Singapore: Pulau Ubin (NHMUK 20130249; USNM 828821). Indonesia: Pantai Cermin, Deli, Sumatra (RMNH 175798); Pulau Dua, Banten, Java (ANSP 225509); Cilcap, Java (RMNH); Madura (RMNH); Lovina, Bali (ZMB 106180); Balikpapan, Kalimantan (AM C. 317961); Makassar, Sulawesi (RNHL 175807; ZMB). Habitat and ecology. This species can be found up to 1.5 m above the ground on trunks of Rhizophora, Nypa and other mangrove trees, in the middle and landward zones of broad mangrove forests in marine and moderately brackish areas. In Thailand, Brandt (1974) noted that it climbs up trees to feed on algae growing on roots and stems; since other Cerithidea species descend the trees to feed on the ground, this observation may be questioned. In Singapore, Berry (1963) observed it on mangrove trees around MHWN and on the mud surface beneath. In Java, Van Benthem Jutting (1956) found it both in mangrove forests and brackish fish ponds, while Mujiono (2009) observed it up to 1.93 m above the ground on mangroves, including Acanthus ilicifolius. Remarks. In molecular phylogenetic analyses, this species forms a clade with C. dohrni, but the two are only reciprocally monophyletic in the COI analysis (Reid et al. 2013; Fig. 1). The uncorrected pairwise distance between them for COI is 0.091, which is low for an interspecific comparison in Cerithidea (only the C. anticipata / reidi comparison, 0.076, is lower). There are, however, small but consistent differences in the shells of the two species, which support their recognition as species. A third species in the C. quoyii group, C. andamanensis, shows differences from these others and characters for their discrimination are listed in Table 1. The ranges of C. quoyii and C. dohrni do not overlap, but those of C. quoyii and C. andamanensis just meet in NE Sumatra (Fig. 11). Over most of its range, C. quoyii is found with C. obtusa (Fig. 8). The latter is a broader, heavier shell with rounded periphery and the animal has a predominantly red, rather than blackish, body (Fig. 2 B, D). Both are used for food in Southeast Asia (Brandt 1974; Hamli et al. 2013)., Published as part of Reid, David G., 2014, The genus Cerithidea Swainson, 1840 (Gastropoda: Potamididae) in the Indo-West Pacific region, pp. 1-65 in Zootaxa 3775 (1) on pages 37-39, DOI: 10.11646/zootaxa.3775.1.1, http://zenodo.org/record/285731, {"references":["Hombron, J. B. & Jacquinot, H. (1848) Atlas d'Histoire Naturelle Zoologie par MM. Hombron et Jacquinot, Chirurgiens de l'Expedition. In: Jacquinot, H. (Ed.), Voyage au Pole Sud et dans l'Oceanie sur les corvettes l'Astrolabe et la Zelee. Livre 26. Gide et Cie, Paris. (Publication date: Clark & Crosnier 2000), pls 17, 21, 23 - 25.","Adams, A. & Reeve, L. (1848 - 1850) Mollusca. In: Adams, A. (Ed.), The Zoology of the Voyage of H. M. S. Samarang. Reeve, Benham & Reeve, London, x + 87 pp.","Lamarck, J. B. P. A. de (1822) Histoire Naturelle de Animaux sans Vertebres. Vols 6 (2), 7. Paris, 232 pp., 711 pp.","Rousseau, L. (1854) Descriptions des mollusques coquilles et zoophytes. In: Hombron, J. B. & Jacquinot, H. (Eds.), Voyage au Pole Sud et dans l'Oceanie sur les corvettes l'Astrolabe et la Zelee … Vol. 5. Zoologie. Gide et J. Baudry, Paris, pp. 1 - 132.","Reid, D. G., Claremont, M., Smith, L., Shamoto, M., Glaubrecht, M. & Ozawa, T. (2013) Mosaics in the mangroves: allopatric diversification of tree-climbing mudwhelks (Gastropoda: Potamididae: Cerithidea) in the Indo-West Pacific. Biological Journal of the Linnean Society, 110, 564 - 580.","Reeve, L. A. (1860) Elements of Conchology. Vol. 1. Reeve, London, 260 pp.","Griffith, E. & Pidgeon, E. (1833 - 1834) The Mollusca and Radiata. In: Griffith, E., (Ed.) The Animal Kingdom arranged in conformity with its organization, by the Baron Cuvier, Vol. 12. Whittaker & Co., London, viii + 601 pp.","Sowerby, G. B. II (1866) Monograph of the genus Cerithidea. In: Reeve, L. A. (Ed.), Conchologia Iconica. Vol. 15. L. A. Reeve, London, pls 1 - 4.","Tapparone-Canefri, C. (1874) Zoologia del viaggio intorno al globo della R. Fregata Magenta. Malacologia. Memorie della Reale Accademia delle Scienze di Torino, Ser. 2, 28, 109 - 265.","Morlet, L. (1889) Catalogue des coquilles recueillis, par M. Pavie, dans le Cambodge et le Royaume de Siam, et description d'especes nouvelles. Journal de Conchyliologie, 37, 121 - 199.","Dautzenberg, P. & Fischer, H. (1906) Liste des mollusques recoltes par M. H. Mansuy en Indo-Chine et au Yunnan et description d'especes nouvelles. Journal de Conchyliologie, 53, 343 - 471.","Van Benthem Jutting, W. S. S. (1956) Systematic studies on the non-marine Mollusca of the Indo-Australian archipelago. V. Critical revision of the Javanese freshwater gastropods. Treubia, 23, 259 - 477.","Poutiers, J. M. (1998) Gastropods. In: Carpenter, K. E. & Niem, V. H. (Eds.), The Living Marine Resources of the Western Central Pacific. Vol. 1. FAO, Rome, pp. 363 - 648.","Swennen, C., Moolenbeek, R. G., Ruttanadakul, N., Hobbelink, H., Dekker, H. & Hajisamae, S. (2001) The molluscs of the southern Gulf of Thailand. Thai Studies in Biodiversity, 4, 1 - 210.","Dharma, B. (2005) Recent & Fossil Indonesian Shells. ConchBooks, Hackenheim, 424 pp.","Reid, D. G., Dyal, P., Lozouet, P., Glaubrecht, M. & Williams, S. T. (2008) Mudwhelks and mangroves: the evolutionary history of an ecological association (Gastropoda: Potamididae). Molecular Phylogenetics and Evolution, 47, 680 - 699. http: // dx. doi. org / 10.1016 / j. ympev. 2008.01.003","Mujiono, N. (2009) Mudwhelks (Gastropoda: Potamididae) from mangroves of Ujung Kulong National Park, Banten. Jurnal Biologi, 13, 51 - 56.","Hamli, H., Idris, M. H., Abu Hena, M. K., Wong, S. K. & Arshad, A. (2013) Checklist and habitat of edible gastropods from Sarawak, Malaysia. Journal of Fisheries and Aquatic Sciences, 8, 412 - 418. http: // dx. doi. org / 10.3923 / jfas. 2013.412.418","Tryon, G. W. (1887) Manual of Conchology. Vol. 9. Tryon, Philadelphia, 488 pp.","Kobelt, W. (1890 b) Eine neue Cerithidea. Nachrichtsblatt der deutschen Malakozoologischen Gesellschaft, 1890, 60 - 61.","Cecalupo, A. (2005) Elenco della famiglia Cerithiidae Ferussac, 1822 (Prosobranchia). Nomenclatura delle specie conosciute o poco note e relativa revisione sistematica. Fossili e Attuali. Vol. I. Quaderni della Civica Stazione Idrobiologica di Milano, 26, 1 - 366.","Martens, E. von (1897 a) Suss- und Brackwasser-Mollusken des Indischen Archipels. In: Weber, M. (Ed.), Zoologische Ergebnisse einer Reise in Niederlandisch Ost-Indien, Vol. 4. E. J. Brill, Leiden, pp. 1 - 331.","Prashad, B. (1921) Report on a collection of Sumatran molluscs from fresh and brackish water. Records of the Indian Museum, 22, 461 - 507.","Brandt, R. A. M. (1974) The non-marine aquatic Mollusca of Thailand. Archiv fur Molluskenkunde, 105, 1 - 423.","Thach, N. N. (2007) Recently Collected Shells of Vietnam. L'Informatore Piceno, Ancona, Italy, 384 pp.","Adams, A. (1855) A monograph of Cerithidea, a genus of Mollusca, with descriptions of several new species, from the collection of High Cuming, Esq,: to which are added, descriptions of two new species of Colina, and one of Donax. Proceedings of the Zoological Society of London, 1854, 83 - 87. http: // dx. doi. org / 10.1111 / j. 1469 - 7998.1854. tb 07236. x","Van Regteren Altena, C. O. (1945) Report upon a collection of Recent shells from Java. Zoologische Mededelingen, 25, 140 - 154.","Ng, P. K. L. & Sivasothi, N. (Eds.) (1999) A Guide to the Mangroves of Singapore II (Animal Diversity). Singapore Science Centre, 168 pp.","Menke, K. T. (1829) Verzeichniss der ansehnlichen Conchylien-Sammlung des Freiherrn von der Malsburg. Pyrmont, vi + 123 pp.","Cecalupo, A. (2006) Elenco della famiglia Cerithiidae Ferussac, 1822 (Prosobranchia). Nomenclatura delle specie conosciute o poco note e relativa revisione sistematica. Attuali. Vol. II. Quaderni della Civica Stazione Idrobiologica di Milano, 27, 1 - 373.","Petit de la Saussaye, M. S. (1851) Description de coquilles nouvelles. Journal de Conchyliologie, 2, 259 - 269.","Berry, A. J. (1963) Faunal zonation in mangrove swamps. Bulletin of the National Museum of Singapore, 32, 90 - 98."]}

Cerithideopsis malayensis new species (Figures 2F���H, N, O, 3B, 4) ? Cerithidea fortunei ��� Casto de Elera, 1896: 329 (not A. Adams, 1855). Cerithidea (Cerithidea) largillierti ��� Hasegawa, 2000: 133 (in part, includes C. largillierti, C. australiensis). Cerithidea largillierti ���Lozouet, 2008: 286, pl. 88, fig. 4 (not Philippi, 1848). Cerithideopsis largillierti ��� Lozouet & Plaziat, 2008: 57, 112, pl. 20, figs 4���6 (in part, includes C. largillierti, C. australiensis). Types. Holotype NHMUK 20100434 /1 (Fig. 2N) and 6 paratypes NHMUK 20100434 /2 (Fig. 2F, O), Kradeh, Kanchanadit, 20 km NE Surat Thani, Thailand. Etymology. From its distribution, centred on the Malay Peninsula. Taxonomic history. The record of ��� Cerithidea fortunei ��� from Mindoro by Casto de Elera (1896) could possibly apply to this species. Also from the Philippines, it has previously been identified and figured as C. largillierti (Lozouet 2008; Lozouet & Plaziat 2008). Diagnosis. Shell small (to 26 mm), brown with one pale spiral band above periphery; animal blackish, yellow edge to foot, 2 yellow bands across snout, yellow tentacles; Malay Peninsula, southern South China Sea, Philippines; COI: HE680619 ��� HE680622, HE680624, HE680625. Material examined. 18 lots. Shell (Fig. 2F���H, N, O): H = 11.0��� 26 mm. Shape elongated conical (H/B = 2.40���2.63, SH = 3.00���3.40); not routinely decollate, 7���11 whorls remaining in well preserved shells, but apex and protoconch always eroded. Spire whorls moderately rounded, suture distinct; spire profile slightly convex, slightly concave towards apex; periphery rounded; relatively delicate. Adult lip only slightly thickened, not flared; sometimes 1 previous lip on final whorl; apertural margin sinuous in side view; anterior canal a weak notch. Sculpture on spire of straight axial ribs, usually becoming curved (opisthocyrt) on last 2���3 whorls, ribs rounded, ribs and interspaces of similar width, 20���27(33) ribs on penultimate whorl, ribs becoming weaker and irregular on final whorl; smallest apical whorls with 3 spiral cords if well preserved, increasing to 6 by whorls 5���6, then becoming weak or disappearing, so that only a trace of 8���10 spiral elements is visible above periphery on penultimate whorl; base with 8���10 cords, of which 2 at periphery slightly enlarged, with marked groove between. No ventrolateral varix, but 0���3 indistinct or prominently raised and rounded varices may be present at any point on last whorl (Fig. 2F). Surface with spiral microstriae on thick periostracum; short periostracal fringe or bristles remaining on larger striae (i.e. 8���10 striae macroscopically visible above periphery) if well preserved; surface sometimes covered by thick black concretion. Colour: brown, with broad cream band above periphery, often concealed by periostracum; band visible by transparency within aperture. Animal (Fig. 3B): sides of foot dark grey, broad yellow margin; head black, snout black with 2 broad transverse yellow stripes (one at tip), tentacles and tentacle bases yellow (examined in two samples from Kedah and Surat Thani). Yellow coloration not preserved after storage in ethanol, but pattern visible as absence of black pigment. Range: Malay Peninsula, southern South China Sea, Philippines. Records: Burma: Kadan Kyun (King I.) (NHMUK 1887.3.10.189). Thailand: Ao Nam Bor, Phuket I. (NHMUK); Kradeh, Kanchanadit, 20 km NE Surat Thani (NHMUK 20100434). Malaysia: Bugang jetty, Merbok estuary, Kedah (NHMUK 20080700); Pulau Lumut, Selangor (USNM 661026). Indonesia: near Bengkalis, Sumatra (T. Goulding & B. Dayrat); Cirebon, Java (NNML); Mempawah, Borneo (MZB). Vietnam: Chilins, Vung Tau (ANSP 330740). Philippines: Abatan R., Bohol (MNHN); Ponson I., Camotes I. (USNM 287982). Habitat and ecology. In Malaysia and Thailand this species was found on firm, peaty mud and in shallow pools with leaf litter, in deep shade at the landward side of the Rhizophora belt. In the Abatan River mangal (Philippines) Lozouet & Plaziat (2008) recorded it on mud in puddles in the shaded inner reaches of the mangrove forest, where it was rare. Remarks. This species is small, usually scarce and difficult to find in its shaded, muddy habitat among leaf litter. It appears to have been overlooked in previous studies of the mangrove-associated molluscs of Southeast Asia (e.g. it was not mentioned by von Martens 1897, Van Benthem Jutting 1956 or Brandt 1974). Molecular samples were available only from the Andaman Sea and Gulf of Thailand coasts of the Malay Peninsula. These showed no phylogeographic structure, in contrast to some other shallow-water marine gastropods in which a genetic break persists between the Indian and Pacific Oceans in this vicinity, consistent with repeated isolation of the two ocean basins during sea-level fluctuations associated with Plio-Pleistocene glaciations (Reid et al. 2006). This could suggest that expansion to the Andaman Sea from the western Pacific (where its sister species C. largillierti occurs) is relatively recent. No molecular samples from the Philippines were available, but this is an area of endemicity in the potamidid genus Cerithidea (Reid et al. 2013). Shells from the Philippines do not differ obviously from those from the rest of the range but, in view of the limited morphological distinction between the three species of the C. largillierti complex, the conspecificity of the population in the Philippines should be tested with molecular data. For comparison between the three species of Cerithideopsis in the Indo-West Pacific, see Remarks on C. australiensis below and Table 2., Published as part of Reid, David G. & Claremont, Martine, 2014, The genus Cerithideopsis Thiele, 1929 (Gastropoda: Potamididae) in the Indo-West Pacific region, pp. 61-80 in Zootaxa 3779 (1) on pages 72-73, DOI: 10.11646/zootaxa.3779.1.8, http://zenodo.org/record/4910133, {"references":["Casto de Elera, R. P. (1896) Catalogo Sistematico de Toda la Fauna de Filipinas. Vol. 3. Moluscos y Radiados. Colegio de Santo Tomas, Manila, 942 + lxiv pp.","Adams, A. (1855) A monograph of Cerithidea, a genus of Mollusca, with descriptions of several new species, from the collection of High Cuming, Esq,: to which are added, descriptions of two new species of Colina, and one of Donax. Proceedings of the Zoological Society of London, 1854, 83 - 87. http: // dx. doi. org / 10.1111 / j. 1469 - 7998.1854. tb 07236. x","Hasegawa, K. (2000) Family Potamididae. In: Okutani, T., (Ed.), Marine Mollusks in Japan. Tokai University Press, Tokyo, pp. 133 - 134.","Philippi, R. A. (1848) Testaceorum novorum centuria (continuatio). Zeitschrift fur Malakozoologie, 1848, 17 - 27.","Lozouet, P. & Plaziat, J. - C. (2008) Mangrove Environments and Molluscs. Abatan River, Bohol and Panglao Islands, Central Philippines. ConchBooks, Hackenheim, 160 pp.","Martens, E. von (1897) Suss- und Brackwasser-Mollusken des Indischen Archipels. In: Weber, M. (Ed.), Zoologische Ergebnisse einer Reise in Niederlandisch Ost-Indien, Vol. 4. E. J. Brill, Leiden, pp. 1 - 331.","Van Benthem Jutting, W. S. S. (1956) Systematic studies on the non-marine Mollusca of the Indo-Australian archipelago. V. Critical revision of the Javanese freshwater gastropods. Treubia, 23, 259 - 477.","Brandt, R. A. M. (1974) The non-marine aquatic Mollusca of Thailand. Archiv fur Molluskenkunde, 105, 1 - 423.","Reid, D. G., Lal, K., Mackenzie-Dodds, J., Kaligis, F., Littlewood, D. T. J. & Williams, S. T. (2006) Comparative phylogeography and species boundaries in Echinolittorina snails in the central Indo-West Pacific. Journal of Biogeography, 33, 990 - 1006. http: // dx. doi. org / 10.1111 / j. 1365 - 2699.2006.01469. x","Reid, D. G., Claremont, M., Smith, L., Shamoto, M., Glaubrecht, M. & Ozawa, T. (2013) Mosaics in the mangroves: allopatric diversification of tree-climbing mudwhelks (Gastropoda: Potamididae: Cerithidea) in the Indo-West Pacific. Biological Journal of the Linnean Society, 110, 564 - 580."]}

Reid, David G., Claremont, Martine (2014): The genus Cerithideopsis Thiele, 1929 (Gastropoda: Potamididae) in the Indo-West Pacific region. Zootaxa 3779 (1): 61-80, DOI: http://dx.doi.org/10.11646/zootaxa.3779.1.8

Cerithideopsis largillierti (Philippi, 1848) (Figures 2A���E, 3A, 4) Cerithium (Potamides) largillierti Philippi, 1848: 20 (China; neotype here designated NHMUK 20130431/1, Shanghai, China, Fig. 2B). Philippi, 1849: 15, Cerithium pl. 1, fig. 7 (cited as fig. 6 in error, p. 15; Fig. 2A herein). Cerithidea (Pirenella) largillierti ���H. Adams & A. Adams, 1854: 293. A. Adams, 1855: 84. Cerithium (Cerithidea) largillierti ���Kobelt, 1890: 65, pl. 13, figs 2, 3 (Cecalupo, 2005: pl. 32, fig. 25). Cerithidea largillierti ��� Dautzenberg & Fischer, 1905: 135. Hirase, 1934: 53, pl. 84, fig. 9. Kuroda & Habe, 1952: 44. Habe & Kosuge, 1967: 34, pl. 13, fig. 12. Habe, 1968: 38, pl. 11, fig. 25. Higo, 1973: 79. Fan, 1989: 38���39, pl. 5, fig. 26. Choe, 1992: 310, pl. 65, fig. 102. Higo & Goto, 1993: 105. Higo, Callomon & Goto, 1999: 87. Fukuda, 2000: 109, 135, figs 5.4c, 5.5c (headfoot). Ma, 2004: 38, pl. 016E. Min et al., 2004: 129, figs 219-1, 219-2. Kojima et al., 2006: 525���535, fig. 2e���g. Li & Huang, 2012: 152, fig. Cerithidea (Cerithidea) largillierti ��� Oyama, 1959: Cerithidea plate, figs 6, 13 (as C. largillerti). Fukuda, 1996: 21, pl. 3, fig. 5. Hasegawa, 2000: 133, pl. 66, fig. 3 (in part, includes C. malayensis, C. australiensis). Hong, Choi & Tsutsumi, 2010: 260, fig. 4a. Fukuda & Kimura, 2012: 28, text fig. a (living animal), b, c (shells). Cerithidea ? largillierti ��� Houbrick, 1984: 16 (in part, includes C. australiensis). Cecalupo, 2005: 318, pl. 32, fig. 25. Cecalupo, 2006: 87, 211. Cerithideopsis largillierti ��� Reid et al., 2008: 680���699 (in part, includes C. australiensis). Lozouet & Plaziat, 2008: 112 (in part, includes C. australiensis, C. malayensis). Cerithidea fortunei A. Adams, 1855: 85 (China; lectotype NHMUK 20130431/1, Fig. 2B; 2 paralectotypes NHMUK 20130431/2, Fig. 2C, 20130413/3, Fig. 2D). Sowerby, 1866: sp. 15, pl. 3, fig. 15a, b. Yen, 1942: 207, pl. 15, fig. 77. Potamides (Cerithidea) fortunei ��� Tryon, 1887: 162, pl. 33, figs 65, 66. Fischer & Dautzenberg, 1904: 416. Iwakawa, 1919: 76. Cerithium montagnei ��� Sowerby, 1855: 888, pl. 186, fig. 286 (in part, includes C. montagnei, C. pulchra; not d���Orbigny, 1841). Cerithidea montagnei ��� Sowerby, 1866: sp. 26 (in part, includes C. montagnei; not d���Orbigny, 1841). Cecalupo, 2006: 70, 202 (in part, includes C. montagnei; not d���Orbigny, 1841). Potamides (Cerithidea) montagnei ��� Tryon, 1887: 161 (in part, includes C. montagnei, C. pulchra; not d���Orbigny, 1841). Cerithidea ornata ��� Yoo, 1995: 61, pl. 9, figs 7, 8. Min et al., 2004: 129, fig. 218 (both not Cerithium ornatum ���A. Adams��� G.B. Sowerby II, 1855 = Cerithidea balteata (A. Adams, 1855)). Taxonomic history. The original description by Philippi (1848) was brief and he only figured the species later (Philippi 1849). This may explain why Adams (1855) listed it as a distinct species in his monograph on Cerithidea, while also naming Cerithidea fortunei. The synonymy of these two names was noted by Sowerby (1855), although he erroneously considered both to be synonyms of Cerithium montagnei (= C. montagnei), a species from the Pacific coast of Central America. This error was pointed out by Carpenter (1857: 343), but Cerithium largillierti was still listed under Cerithidea montagnei by Sowerby (1866) and Tryon (1887). Meanwhile, Cerithidea fortunei continued to be widely used as the name of the present species (Sowerby 1866; Tryon 1887; Fischer & Dautzenberg 1904; Iwakawa 1919). The name Cerithidea largillierti was reinstated by Dautzenberg & Fischer (1905) and thereafter this specific name has been almost universally used. There has been occasional misidentification as ��� Cerithidea ornata ��� (Yoo 1995; Min et al. 2004). No type material of Cerithium largillierti has been traced (not present in MNHN, ZMB or SMF, which all contain some specimens of other species collected by Largilliert and/or described by Philippi). Philippi���s (1849) figure could serve as lectotype, but is not clearly drawn (Fig. 2A) and an actual specimen is desirable. Therefore, a shell from Shanghai is here designated the neotype (NHMUK 20130431/1; Fig. 2B). This shell is also one of the syntypes of C. fortunei A. Adams, 1855. To avoid any possible confusion, and to reinforce the synonymy, this specimen is also designated the lectotype of C. fortunei. Diagnosis. Shell of medium size (to 36 mm), brown with two pale spiral bands above periphery; animal black with yellow tip to snout and tentacle bases; northern Vietnam, China, Korea, southern Japan; COI: AM 932788, AM 932789, HE680618, HE680623. Material examined. 22 lots. Shell (Fig. 2A���E): H = 23.9���35.9 mm. Shape elongated conical (H/B = 2.17���2.42, SH = 2.88���3.31); not routinely decollate, 9���11 whorls remaining in well preserved shells, but apex and protoconch always eroded. Spire whorls rounded, suture distinct; spire profile slightly convex, slightly concave towards apex; periphery rounded; relatively solid. Adult lip only slightly thickened, not flared; sometimes 1���2 previous lips on final whorl; apertural margin sinuous in side view; anterior canal a weak notch. Sculpture on spire of straight axial ribs, usually becoming curved (opisthocyrt) on last 2���3 whorls, ribs rounded, ribs and interspaces of similar width, 20���26 ribs on penultimate whorl, ribs becoming weaker and irregular on final whorl; 3���6 spiral cords on smallest apical whorls if well preserved, macroscopic spiral sculpture usually absent above periphery of larger whorls, sometimes 10���13 weak spiral elements visible between axial ribs on last 2 whorls; base with 8���9 cords, of which two at periphery are strong, with marked groove between, those towards columella becoming smaller. No ventrolateral varix, but 0���2 indistinct or prominently raised and rounded varices may be present at any point on last 2 whorls (Fig. 2D). Surface with spiral microstriae on thick periostracum, short periostracal fringe or bristles remaining on larger striae (i.e. 10���13 striae macroscopically visible above periphery) if well preserved; shiny where periostracum worn away; surface often covered by thick black concretion, presumably of microscopic algae and sediment. Colour: brown, with pale band just above periphery and another at shoulder, often indistinct beneath periostracum; aperture brown with indistinct pale bands. Animal (Fig. 3A): sides of foot black, paler towards margin; head grey, snout black, snout tip and tentacle bases yellow. Yellow coloration not preserved after storage in ethanol, but pattern visible as absence of black pigment (examined in two lots from Japan and Korea). Range: N Vietnam, China, Taiwan, S Korea, S Japan. Records: Vietnam: Quikim, Hai Phong (NHMUK; ZMSP). China: Tsim Bei Tsui, Hong Kong (NHMUK); Shanghai (NHMUK 20130431; ZMB; ANSP 275721); ���throughout Chinese coasts��� (Ma 2004: 38). Taiwan: Hsinchu (Yan Yu-Xiu; photo seen). S Korea: Daehang-ri, Byeonsan-myeon, Buan-gun, Jeollabuk-do (NHMUK 20070408); Suncheon Bay (Kamimura et al. 2010). Japan: Nagasaki (USNM 343900); Hikawa, Kumamoto Pref. (NHMUK); Onoda City, Yamaguchi Pref. (NHMUK); Tokyo (USNM 343901; Callomon & Goto 1999: quoting Okamoto 1995; Hasegawa 2000). Habitat and ecology. Mud flats at high tidal levels, on bare mud and among Phragmites reeds, often at river mouths; among saltmarsh vegetation (Korea) and at margins of brackish shrimp ponds (northern Vietnam). Remarks. Over much of its range this species overlaps with two other potamidids, Cerithidea sinensis (Philippi) (China and northern Vietnam) and Cerithidea tonkiniana Mabille (Vietnam to southern Japan) (see Reid 2014 for detailed descriptions). The shells of these Cerithidea species are decollate as adults, the aperture is more strongly flared (and planar, not sinuous, in Cerithidea tonkiniana), the whorls less rounded, the axial ribs less numerous (14���19 in Cerithidea sinensis; 11���25 in Cerithidea tonkiniana), the periostracum is thin and the ventrolateral varix often present. The living animals also differ. That of Cerithidea sinensis has a cream head and foot with grey mottling, the tip of the snout is yellow with two transverse black bands just behind, and the tentacle yellow at the base with a black stripe across the eye. That of Cerithidea tonkiniana has a blackish foot, a grey head with cream spots, the anterior half of the snout is blackish with yellow spots arranged in 2���3 transverse bands, and the tentacle is grey with black rings and a black base. For comparison between the three species of Cerithideopsis in the Indo-West Pacific, see Remarks on C. australiensis below and Table 2. There have been two genetic studies of C. largillierti. Kojima et al. (2006) found two shared haplotypes of COI and no significant differences in frequencies between Japan and China. This was interpreted as suggesting recent migration between the two regions, either during the last glacial stage when there was a connection between Kyushu and Korea, or through a possible planktotrophic larva (larval development is unknown in this species). Kamimura et al. (2010) reported that COI haplotypes were of low diversity in Suncheon Bay, South Korea. The species inhabits threatened wetlands in southern Japan, where it has been listed as endangered (Fukuda 1996) or vulnerable (Fukuda & Kimura 2012)., Published as part of Reid, David G. & Claremont, Martine, 2014, The genus Cerithideopsis Thiele, 1929 (Gastropoda: Potamididae) in the Indo-West Pacific region, pp. 61-80 in Zootaxa 3779 (1) on pages 68-70, DOI: 10.11646/zootaxa.3779.1.8, http://zenodo.org/record/4910133, {"references":["Philippi, R. A. (1848) Testaceorum novorum centuria (continuatio). Zeitschrift fur Malakozoologie, 1848, 17 - 27.","Philippi, R. A. (1849) Abbildungen und Beschreibungen neuer oder wenig gekannter Conchylien. Vol. 3. Part 4. Theodor Fischer, Cassel, 28 pp.","Adams, A. (1855) A monograph of Cerithidea, a genus of Mollusca, with descriptions of several new species, from the collection of High Cuming, Esq,: to which are added, descriptions of two new species of Colina, and one of Donax. Proceedings of the Zoological Society of London, 1854, 83 - 87. http: // dx. doi. org / 10.1111 / j. 1469 - 7998.1854. tb 07236. x","Cecalupo, A. (2005) Elenco della famiglia Cerithiidae Ferussac, 1822 (Prosobranchia). Nomenclatura delle specie conosciute o poco note e relativa revisione sistematica. Fossili e Attuali. Vol. I. Quaderni della Civica Stazione Idrobiologica di Milano, 26, 1 - 366.","Dautzenberg, P. & Fischer, H. (1905) Liste des mollusques recoltes par M. le Capitaine de Fregate Blaise au Tonkin, et description d'especes nouvelles. Journal de Conchyliologie, 53, 85 - 234. http: // dx. doi. org / 10.5962 / bhl. title. 13158","Hirase, S. (1934) A Collection of Japanese Shells with Illustrations in Natural Colours. M. Sanshodo, Tokyo, 217 pp.","Kuroda, T. & Habe, T. (1952) Check List and Bibliography of the Recent Marine Mollusca of Japan. L. W. Stach, Tokyo, 210 pp.","Habe, T. & Kosuge, S. (1967) The Standard Book of Japanese Shells in Colour, Vol. III. Hoikusha, Osaka, xviii + 223 pp.","Habe, T. (1968) Shells of the Western Pacific in Color. Vol. 2. 2 nd ed. Hoikusha, Osaka, 233 pp.","Higo, S. (1973) A Catalogue of Molluscan Fauna of the Japanese Islands and the Adjacent Area. Isahaya, 397 + 61 pp.","Fan, C. (Ed.) (1989) Mollusca of Huanghai and Bohai. Nongye Chubanshe, Agricultural Publishing House, Beijing, 309 pp.","Choe, B. L. (1992) Mollusca (II). Illustrated Encyclopaedia of Fauna and Flora of Korea. Vol. 33. Ministry of Education, Republic of Korea, 860 pp.","Higo, S. & Goto, Y. (1993) A Systematic List of Molluscan Shells from the Japanese Islands and the Adjacent Areas. Eru Malacological Publisher, Osaka, 3 + 22 + 693 + 13 + 149 pp.","Higo, S., Callomon, P. & Goto, Y. (1999) Catalogue and Bibliography of the Marine Shell-Bearing Mollusca of Japan. Elle Scientific Publications, Osaka, 749 pp.","Fukuda, H. (2000) Gastropods I: general view. In: Sato, M. (Ed.) Life in Ariake Sea: Biodiversity in Tidal Flats and Estuaries. Kaiyu-sha, Tokyo, pp. 213 - 252.","Ma, X. (2004) Order Mesogastropoda. In: Qi, Z. (Ed.), Seashells of China. China Ocean Press, Beijing, pp. 31 - 81.","Min, D. - K., Lee, J. - S., Koh, D. - B. & Je, J. - G. (2004) Mollusks in Korea. Min Molluscan Research Institute, Seoul, 566 pp.","Kojima, S., Kamimura, S., Iijima, A., Kimura, T., Kurozumi, T. & Furota, T. (2006) Molecular phylogeny and population structure of tideland snails in the genus Cerithidea around Japan. Marine Biology, 149, 525 - 535. http: // dx. doi. org / 10.1007 / s 00227 - 005 - 0183 - 2","Li, R. & Huang, Z. (2012) Class Gastropoda. In: Huang, Z. & Liu, M. (Eds.), The Living Species and their Illustrations in China's Seas (Part II). An Illustrated Guide to Species in China's Sea. Vol. 4. Ocean Press, Beijing, pp. 102 - 279.","Oyama, K. (1959) Cerithidea. In: Oyama, K. (Ed.), The Molluscan Shells. Vol. 2. Resources Exploitation Institute, Tokyo, unpaginated.","Fukuda, H. (1996) Gastropoda. In: Hanawa, S. & Sakuma, H. (Eds.), Present Status of Estuarine Locales and Benthic Invertebrates Occurring in Estuarine Environment in Japan. WWF Japan Science Report, 3, 11 - 63.","Hasegawa, K. (2000) Family Potamididae. In: Okutani, T., (Ed.), Marine Mollusks in Japan. Tokai University Press, Tokyo, pp. 133 - 134.","Hong, J. - S., Choi, J. - W. & Tsutsumi, H. (2010) Concluding remarks on the joint survey of macrobenthic fauna on Suncheon Tidal Flats by the participants of \" Korea and Japan Joint Symposium on Biology of Tidal Flats 2009 \". Plankton & Benthos Research, 5 (suppl.), 255 - 263. http: // dx. doi. org / 10.3800 / pbr. 5.255","Fukuda, H. & Kimura, S. (2012) Cerithidea (Cerithidea) largillierti (Philippi, 1848). In: Japanese Association of Benthology. (Ed.), Threatened Animals of Japanese Tidal Flats: Red Data Book of Seashore Benthos. Tokai University Press, Hadano, pp. 28.","Houbrick, R. S. (1984) Revision of higher taxa in genus Cerithidea (Mesogastropoda: Potamididae) based on comparative morphology and biological data. American Malacological Bulletin, 2, 1 - 20.","Cecalupo, A. (2006) Elenco della famiglia Cerithiidae Ferussac, 1822 (Prosobranchia). Nomenclatura delle specie conosciute o poco note e relativa revisione sistematica. Attuali. Vol. II. Quaderni della Civica Stazione Idrobiologica di Milano, 27, 1 - 373.","Reid, D. G., Dyal, P., Lozouet, P., Glaubrecht, M. & Williams, S. T. (2008) Mudwhelks and mangroves: the evolutionary history of an ecological association (Gastropoda: Potamididae). Molecular Phylogenetics and Evolution, 47, 680 - 699. http: // dx. doi. org / 10.1016 / j. ympev. 2008.01.003","Lozouet, P. & Plaziat, J. - C. (2008) Mangrove Environments and Molluscs. Abatan River, Bohol and Panglao Islands, Central Philippines. ConchBooks, Hackenheim, 160 pp.","Sowerby, G. B. II (1866) Monograph of the genus Cerithidea. In: Reeve, L. A. (Ed.), Conchologia Iconica. Vol. 15. L. A. Reeve, London, pls 1 - 4.","Yen, T-C. (1942) A review of Chinese gastropods in the British Museum. Proceedings of the Malacological Society of London, 24, 170 - 289.","Tryon, G. W. (1887) Manual of Conchology. Vol. 9. Tryon, Philadelphia, 488 pp.","Fischer, H. & Dautzenberg, P. (1904) Catalogue des mollusqes terrestres et fluviatiles de l'Indo-Chine orientale cites jusqu'a ce jour. In: Pavie, A. (Ed.), Mission Pavie Indo-Chine 1879 - 1895. Etudes Diverses. III. Recherches sur l'Histoire Naturelle de l'Indo-Chine Orientale. E. Leroux, Paris, pp. 390 - 450.","Iwakawa, T. (1919) Catalogue of Japanese Mollusca in the Natural History Department, Tokyo Imperial Museum. Tokyo Imperial Museum, Tokyo, 8 + 318 + 5 + 95 + 39 pp. http: // dx. doi. org / 10.5962 / bhl. title. 13147","Sowerby, G. B. II (1855) Monograph of the genus Cerithium, Adanson. In: Sowerby, G. B. II (Ed.), Thesaurus Conchyliorum, or Figures and Descriptions of Recent Shells. Vol. 2. part 16. Sowerby, London, pp. 847 - 899.","Yoo, J. - S. (1995) Korean Shells in Colour. Il Ji Sa Publishing Co., Seoul, 196 pp.","Carpenter, P. P. (1857) Catalogue of the Collection of Mazatlan Shells in the British Museum: Collected by Frederick Reigen. British Museum, London, xvi + 552 pp.","Kamimura, S., Itoh, H., Ozeki, S. & Kojima, S. (2010) Molecular diversity of Cerithidea gastropods inhabiting Suncheon Bay, and the Japanese and Ryukyu Islands. Plankton & Benthos Research, 5 (suppl.), 250 - 254. http: // dx. doi. org / 10.3800 / pbr. 5.250","Reid, D. G. (2014) The genus Cerithidea Swainson, 1840 (Gastropoda: Potamididae) in the Indo-West Pacific region. Zootaxa, 3775 (1), 1 - 65. http: // dx. doi. org / 10.11646 / zootaxa. 3775.1.1"]}

Cerithideopsis australiensis new species (Figures 2I���M, P���T, 3C, D, 4) Cerithidea largillierti ��� Hedley, 1909: 357. Hinton, 1978: pl. 7, fig. 1. Hinton, 1979: pl. 6, fig. 1. Healy, 1983: 57���75, figs 50��� 53, 55, 63, 69, 95���98 (euspermatozoa). Wells & Bryce, 1986: 55, pl. 11, fig. 129. Jarrett, 2011: 40, fig. 96 (all not Philippi, 1848). Cerithidea ? largillierti ��� Houbrick, 1984: 16 (in part, includes C. largillierti). Cerithidea (Cerithidea) largillierti ��� Wilson, 1993: 133, pl. 15, fig. 12 (not Philippi, 1848). Hasegawa, 2000: 133 (in part, includes C. largillierti, C. malayensis). Cerithideopsis largillierti ��� Reid et al., 2008: 680���699, fig. 2 (phylogeny and shell; in part, includes C. largillierti). Lozouet & Plaziat, 2008: 112 (in part, includes C. largillierti, C. malayensis). Willan, 2013: 78, fig. 9 (not Philippi, 1848). Types. Holotype AM C.478294 (Fig. 2L, M) and 2 paratypes NHMUK 20130435, Cockle Bay, Magnetic Island, Queensland, Australia. Etymology. From its distribution, largely confined to Australia. Taxonomic history. Previously, this species has always been known under the specific name largillierti. Reid et al. (2008) first suggested that C. largillierti s. l. consisted of two species, from Japan and Australia respectively, with a relatively high genetic (K2P) distance for COI of 15%. Diagnosis. Shell relatively large (to 48 mm), brown with 1���2 pale spiral bands, or white with 1���3 brown bands; animal black with dense yellow spots; tropical Australia and Gulf of Papua; COI: AM 932790, AM 932791, HE680615 ��� HE680617. Material examined. 42 lots. Shell (Fig. 2I���M, P���T): H = 15.2���48.2 mm. Shape elongated conical (H/B = 2.30���2.64, SH = 3.13���3.48); not routinely decollate, 10���14 whorls remaining in well-preserved shells, but apex and protoconch always eroded. Spire whorls rounded, suture distinct; spire profile very slightly convex, slightly concave towards apex; periphery rounded; relatively solid. Adult lip only slightly thickened and flared; sometimes up to 3 previous lips and strong growth lines near end of final whorl; apertural margin sinuous in side view; anterior canal a weak notch. Sculpture on spire of straight axial ribs, becoming curved (opisthocyrt) on last 2���3 whorls, ribs rounded, ribs and interspaces of similar width, 15���24(29) ribs on penultimate whorl, ribs becoming weaker and irregular on final whorl; smallest apical whorls with 2���3 spiral cords if well preserved, increasing to 5 by whorls 5���6, then disappearing, so that only a trace of 8���10 spiral elements is visible above periphery on penultimate whorl, stronger posteriorly; base with 8��� 10 cords, of which 2 at periphery slightly enlarged, with marked groove between. No ventrolateral varix, but 0���3 indistinct or prominently raised and rounded varices may be present at any point on last whorl, up to 7 in total on last 3 whorls (Fig. 2J, M, Q, S). Surface with spiral microstriae on thick periostracum (Fig. 3D); short periostracal fringe or bristles remain on larger striae (i.e. 8���10 striae macroscopically visible above periphery) if well preserved. Colour: brown, with broad cream band above periphery, sometimes additional pale band at suture; pattern often concealed by periostracum; in northwestern Australia shells are white with 1���2 brown bands above periphery and sometimes a brown line between 2 strong outermost basal grooves (Fig. 2R���T); bands visible by transparency within aperture. Animal (Fig. 3C): sides of foot and snout black with dense yellow spots, 2 larger yellow spots near tip of snout; tentacle bases yellow (Fig. 3C from Northern Territory). Yellow coloration not preserved after storage in ethanol, but pattern visible as absence of black pigment (examined in two lots from Proserpine and Thirsty Sound, Queensland). Range: Tropical Australia and Gulf of Papua. Records: Western Australia: Cossack (AM C.439257); Lookout Hill, Broome (NHMUK 20130437); Buccaneer Archipelago (AM C.042509). Northern Territory: Ludmilla Creek, Darwin (NHMUK); Bing Bong station (AM C.412930). Queensland: Forsyth I. (AM C.108447); Mapoon (AM C.014282); Missionary Bay, Hinchinbrook I. (NHMUK); Cockle Bay, Magnetic I. (NHMUK 20130435); Plum Tree, Thirsty Sound (AM C.419622); Russell I., Moreton Bay (AM C.410423); Lota Creek, Waterloo Bay, SW Moreton Bay (NTM P.49902). Papua New Guinea: 30 miles E Port Moresby (AM C.82943). Wells & Bryce (1986: 55) recorded this species from ���North West Cape, W.A. to Queensland ���. The southernmost records are from Moreton Bay, southern Queensland. Hinton (1979: caption of pl. 7) observed that this species was ���reasonably common in Gulf of Papua ���. Habitat and ecology. In northern Queensland it is found on mud and in shallow pools with leaf litter, in the shade of Ceriops trees on the edges of saltpans in mangrove areas. The snails bury themselves in the mud during neap tides. In Broome, Western Australia, it was found on soft mud beneath Avicennia trees in a wide belt of this tree fringing a coastal mudflat. Willan (2013) reported that it inhabits the middle and lower tidal zones of mangrove forests where the mud is soft, and remains in pools when the tide recedes. Remarks. Shells from Western Australia are always pale in colour and of unusually thick texture (Fig. 2R���T). This marine region is known for its high endemicity and the mangrove-associated gastropods include the endemic Cerithidea reidi Houbrick (Reid 2014), Littoraria cingulata (Philippi) and L. sulculosa (Philippi), besides several distinctive regional forms with low genetic divergence (Reid 1986; Reid, Dyal & Williams 2010). White shells with brown lines (2 above periphery and 2 below) are occasionally found as far east as Darwin (NTM P.009694). Genetic study is required to test the status of the western form of C. australiensis. The three species of Cerithideopsis in the Indo-West Pacific are extremely similar in shell characters and there are no entirely diagnostic characters by which to separate them (Table 2). In overall size, C. malayensis is typically small (not exceeding 26.0 mm in shell length), while C. australiensis reaches largest size (up to 48.2 mm in shell length). Their shapes and rib counts show broad overlap. In shell colour, C. largillierti typically has two pale bands above the periphery, while C. malayensis has one and colour is more variable in C. australiensis, but this feature is not useful when the shell is encrusted with mud and algae, covered by a thick periostracum or when severely eroded. The headfoot of C. largillierti is black with yellow tentacle bases and tip of snout when living; that of C. malayensis is black with two yellow stripes across the snout and a yellow edge to the foot, while that of C. australiensis is black with yellow spots. In ethanol-preserved specimens the yellow coloration fades almost to white, but the black pigmentation remains. Available information indicates that these colour patterns are speciesspecific, but this requires confirmation. The distributions of the three species are entirely allopatric (Fig. 4) and this provides the simplest means of identification. Cerithideopsis australiensis is of small size relative to edible potamidids such as species of Terebralia, Telescopium and Cerithidea, and has not been found in densities that would appear sufficient to warrant collection for food. Willan (2013) noted that C. australiensis is never consumed by Aboriginal people in Australia. However, this has not always been the case; the species dominates many middens dated to around 2000���2500 BP on the coast of Papua New Guinea southeast of Port Moresby (Petchey et al 2012; specimens from Caution Bay examined)., Published as part of Reid, David G. & Claremont, Martine, 2014, The genus Cerithideopsis Thiele, 1929 (Gastropoda: Potamididae) in the Indo-West Pacific region, pp. 61-80 in Zootaxa 3779 (1) on pages 73-75, DOI: 10.11646/zootaxa.3779.1.8, http://zenodo.org/record/4910133, {"references":["Hedley, C. (1909) The marine fauna of Queensland. Proceedings of the Australasian Association for the Advancement of Science, Section D Biology, 1909, 329 - 371.","Hinton, A. G. (1978) Guide to Australian Shells. Robert Brown, Port Moresby, 82 pp.","Hinton, A. G. (1979) Guide to Shells of Papua New Guinea. Robert Brown, Port Moresby, 68 pp.","Healy, J. M. (1983) Ultrastructure of euspermatozoa of cerithiacean gastropods (Prosobranchia: Mesogastropoda). Journal of Morphology, 178, 57 - 75. http: // dx. doi. org / 10.1002 / jmor. 1051780105","Wells, F. E. & Bryce, C. W. (1986) Seashells of Western Australia. Western Australian Museum, Perth, 207 pp.","Jarrett, A. G. (2011) Marine Shells of the Whitsunday Coast & Islands. Alan G. Jarrett, Cannonvale, 268 pp.","Philippi, R. A. (1848) Testaceorum novorum centuria (continuatio). Zeitschrift fur Malakozoologie, 1848, 17 - 27.","Houbrick, R. S. (1984) Revision of higher taxa in genus Cerithidea (Mesogastropoda: Potamididae) based on comparative morphology and biological data. American Malacological Bulletin, 2, 1 - 20.","Wilson, B. R. (1993) Australian marine shells. Prosobranch gastropods. Part I. Odyssey Publishing, Kallaroo, Western Australia, 407 pp.","Hasegawa, K. (2000) Family Potamididae. In: Okutani, T., (Ed.), Marine Mollusks in Japan. Tokai University Press, Tokyo, pp. 133 - 134.","Reid, D. G., Dyal, P., Lozouet, P., Glaubrecht, M. & Williams, S. T. (2008) Mudwhelks and mangroves: the evolutionary history of an ecological association (Gastropoda: Potamididae). Molecular Phylogenetics and Evolution, 47, 680 - 699. http: // dx. doi. org / 10.1016 / j. ympev. 2008.01.003","Lozouet, P. & Plaziat, J. - C. (2008) Mangrove Environments and Molluscs. Abatan River, Bohol and Panglao Islands, Central Philippines. ConchBooks, Hackenheim, 160 pp.","Willan, R. C. (2013) A key to the potamidid snails (longbums, mudcreepers and treecreepers) of northern Australia. Northern Territory Naturalist, 24, 68 - 80.","Reid, D. G. (2014) The genus Cerithidea Swainson, 1840 (Gastropoda: Potamididae) in the Indo-West Pacific region. Zootaxa, 3775 (1), 1 - 65. http: // dx. doi. org / 10.11646 / zootaxa. 3775.1.1","Reid, D. G. (1986) The Littorinid Molluscs of Mangrove Forests in the Indo-Pacific Region: the genus Littoraria. British Museum (Natural History), London, xv + 228 pp.","Reid, D. G., Dyal, P. & Williams, S. T. (2010) Global diversification of mangrove fauna: a molecular phylogeny of Littoraria (Gastropoda: Littorinidae). Molecular Phylogenetics and Evolution, 55, 185 - 201. http: // dx. doi. org / 10.1016 / j. ympev. 2009.09.036","Petchey, F., Ulm, S., David, B., McNiven, I. J., Asmussen, B., Tomkins, H., Richards, T., Rowe, C., Leavesley, M., Mandui, H. & Stanisic, J. (2012) 14 C marine reservoir variability in herbivores and deposit-feeding gastropods from an open coastline, Papua New Guinea. Radiocarbon, 54, 967 - 978."]}

Genus Cerithideopsis Thiele, 1929 Cerithidea (Cerithideopsis) Thiele, 1929: 206 (type species by original designation Cerithidea iostoma (Pfeiffer) = Potamides iostomus Pfeiffer, 1839 = C. pliculosa (Menke, 1829)). Taxonomic history. Thiele (1929) introduced Cerithideopsis as a subgenus of Cerithidea on the basis of the short, rather square, rachidian tooth and other small differences from the radula of Cerithidea decollata (L.). As originally conceived, the new subgenus included species now assigned to the genus Cerithideopsilla (itself described as a new section of the subgenus by the same author, on the basis of its shell characters and IWP distribution). Pirenella was retained as a distinct genus, for forms of P. conica (Blainville). In his original sense, Thiele���s (1929) nominate section Cerithideopsis included only American species. Subsequently, Cerithideopsis was generally treated as a purely American subgenus of Cerithidea (Wenz 1938; Bequaert 1942; Houbrick 1984). As a result of the molecular phylogenetic analysis by Reid et al. (2008), Cerithideopsis was raised to generic rank and the IWP species of the C. largillierti complex were added. Diagnosis. Shell: moderately large (to 48 mm), delicate to solid, elongate, spire not decollate; sculpture of axial ribs, weakening on body whorl; 1���2 swollen varices may be present anywhere on body whorl (not routinely in a ventrolateral position) and others scattered over spire whorls; spiral ridges present on base, outermost 1���2 strongest with deep groove between, ridges absent or sometimes present above periphery; aperture circular, peristome sometimes flared and thickened in adult, planar or sinuous in side view, with weak anterior canal and slight posterior groove; columella straight or very slightly twisted, lacking folds; colour dark with peripheral white band and sometimes other spiral lines. Radula: rachidian tooth broad, square to pentagonal, with long central cusp and 2 denticles on each side; lateral tooth with long lateral extension; outer marginal with numerous small cusps and wide flange on outer side of basal shaft (Bright 1958: C. californica; Houbrick 1984: C. scalariformis). Remarks. The comparative morphology of Cerithideopsis, Cerithidea and Cerithideopsilla has been described in detail by Houbrick (1984). Their respective monophyly was established in the molecular study by Reid et al. (2008), who also gave brief morphological diagnoses. A more detailed taxonomic account of Cerithidea was given by Reid (2014). The three western Atlantic species of Cerithideopsis are C. pliculosa, C. costata and C. scalariformis; these are relatively well known (Bequaert 1942; Abbott 1974). Traditionally, six species have been accepted on the American west coast (Keen 1971; Abbott 1974), but a recent molecular study (Miura et al. 2010) has suggested that this should be reduced to three: C. californica, C. montagnei and C. pulchra (C.B. Adams, 1852). Molecular discrimination of the American species is complicated by two cases of dispersal across the Panamanian Isthmus and subsequent introgression of mitochondrial haplotypes (Miura et al. 2012). The six Cerithideopsis species from the eastern Pacific and western Atlantic, and the three from the IWP described here, form two sister clades (Reid et al. 2008; Miura et al. 2010; Fig. 1). Several morphological characters are restricted to the IWP species, including the thick periostracum, the absence of a strongly thickened and flared peristome, and the yellow markings of the living animal���contrasting with the brown to black animals described by Bright (1958) in C. californica and Houbrick (1984) in C. scalariformis. The deeper phylogeny of the family (Reid et al. 2008) suggests that these are likely synapomorphies. Spawn and larval development have been reported in several species. The American Cerithideopsis species lay strings of eggs on mangrove roots, fallen leaves, algae or the mud surface (Houbrick 1984; Miura, Frankel & Torchin 2011). In the western Atlantic, C. scalariformis (Houbrick 1984) and C. pliculosa (Miura et al. 2011) hatch as late swimming-crawling veligers without a planktonic stage. In the eastern Pacific, C. californica in California either settles immediately upon hatching (Miura et al. 2011, quoting unpublished thesis by MacDonald 1967) or emerges at the crawling stage (Race 1981). However, in Panama C. californica shows a planktonic phase of up to 19 days (Miura et al. 2011). Development of the IWP species has not been recorded. The eastern Pacific and western Atlantic species of Cerithideopsis are all found at high tidal levels on sheltered sedimentary shores, often in estuaries and lagoons. They frequently shelter beneath mangroves and salt marsh plants, and all have been recorded to climb on vegetation at least occasionally, particularly to avoid submersion during high tide (Houbrick 1984; Cantera, Thomassin & Arnaud 1999; D.G. Reid unpublished observation of C. montagnei). However, they do not appear to display the regular climbing on mangroves and other emergent vegetation (involving avoidance of submergence and downward migration to feed) that is characteristic of Cerithidea species (Reid 2014). Climbing behaviour appears to have been independently derived in these two genera (Reid et al. 2008). Burrowing into the mud during cold spells has been recorded in C. californica (Race 1981). The IWP Cerithideopsis species have never been seen to climb on vegetation and appear to have no aversion to being submerged in shallow pools; C. australiensis is known to bury in the mud during neap tides (see below)., Published as part of Reid, David G. & Claremont, Martine, 2014, The genus Cerithideopsis Thiele, 1929 (Gastropoda: Potamididae) in the Indo-West Pacific region, pp. 61-80 in Zootaxa 3779 (1) on pages 67-68, DOI: 10.11646/zootaxa.3779.1.8, http://zenodo.org/record/4910133, {"references":["Thiele, J. (1929, 1931) Handbuch der Systematischen Weichtierkunde. Parts 1, 2. Gustav Fischer, Jena, 777 pp.","Pfeiffer, L. (1839) Bericht uber die Ergebnisse meiner Reise nach Cuba im Winter 1838 - 1839. Archiv Fur Naturgeschichte, 5, 346 - 358.","Menke, K. T. (1829) Verzeichniss der ansehnlichen Conchylien-Sammlung des Freiherrn von der Malsburg. Pyrmont, vi + 123 pp.","Wenz, W. (1938) Gastropoda. Teil I: Allgemeiner Teil und Prosobranchia. In: Schindewolf, O. H. (Ed.), Handbuch der Palaozoologie. Vol. 6. Borntraeger, Berlin, xii + 948 pp.","Bequaert, J. C. (1942) Cerithidea and Batillaria in the western Atlantic. Johnsonia, 5, 1 - 11.","Houbrick, R. S. (1984) Revision of higher taxa in genus Cerithidea (Mesogastropoda: Potamididae) based on comparative morphology and biological data. American Malacological Bulletin, 2, 1 - 20.","Reid, D. G., Dyal, P., Lozouet, P., Glaubrecht, M. & Williams, S. T. (2008) Mudwhelks and mangroves: the evolutionary history of an ecological association (Gastropoda: Potamididae). Molecular Phylogenetics and Evolution, 47, 680 - 699. http: // dx. doi. org / 10.1016 / j. ympev. 2008.01.003","Bright, D. B. (1958) Morphology of the common mudflat snail, Cerithidea californica. Bulletin of the Southern California Academy of Sciences, 57, 127 - 139.","Reid, D. G. (2014) The genus Cerithidea Swainson, 1840 (Gastropoda: Potamididae) in the Indo-West Pacific region. Zootaxa, 3775 (1), 1 - 65. http: // dx. doi. org / 10.11646 / zootaxa. 3775.1.1","Abbott, R. T. (1974) American Seashells. 2 nd Edition. Van Nostrand Reinhold, New York, 663 pp.","Keen, A. M. (1971) Sea Shells of Tropical West America. Ed. 2. Stanford University Press, Stanford, xiv + 1064 pp.","Miura, O., Torchin, M. E., & Bermingham, E. (2010) Molecular phylogenetics reveals differential divergence of coastal snails separated by the Isthmus of Panama. Molecular Phylogenetics and Evolution, 56, 40 - 48. http: // dx. doi. org / 10.1016 / j. ympev. 2010.04.012","Miura, O., Torchin, M. E., Bermingham, E., Jacobs, D. K. & Hechinger, R. F. (2012) Flying shells: historical dispersal of marine snails across Central America. Proceedings of the Royal Society B, 279, 1061 - 1067. http: // dx. doi. org / 10.1098 / rspb. 2011.1599","Miura, O., Frankel, V. & Torchin, M. E. (2011) Different developmental strategies in geminate snails, Cerithideopsis californica and C. pliculosa, across the Isthmus of Panama. Journal of Molluscan Studies, 77, 255 - 258. http: // dx. doi. org / 10.1093 / mollus / eyr 012","Race, M. S. (1981) Field ecology and natural history of Cerithidea californica (Gastropoda: Prosobranchia) in San Francisco Bay. Veliger, 24, 18 - 27.","Cantera, J. R., Thomassin, B. A. & Arnaud, P. M. (1999) Faunal zonation and assemblages in the Pacific Colombian mangroves. Hydrobiologia, 413, 17 - 33. http: // dx. doi. org / 10.1007 / 978 - 94 - 011 - 4078 - 2 _ 3"]}

SUBFAMILY LACUNINAE GRAY, 1857 Risellidae Kesteven, 1903: 621–631. Cremnoconchinae Preston, 1915: 64. Bembiciidae Finlay, 1928: 241. 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., Published as part of 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), pp. 93-135 in Zoological Journal of the Linnean Society 167 (1) on page 98, DOI: 10.1111/j.1096-3642.2012.00875.x, http://zenodo.org/record/5292273, {"references":["Gray JE. 1857. Guide to the systematic distribution of Mollusca in the British Museum. Part I. London: British Museum.","Kesteven HL. 1903. Notes on Prosobranchiata, no. 2. - Littorinacea. Proceedings of the Linnean Society of New South Wales. 27: 620 - 636.","Preston HB. 1915. The fauna of British India: Mollusca (freshwater Gastropoda and Pelecypoda). London: Taylor & Francis.","Finlay HJ. 1928. The Recent Mollusca of the Chatham Islands. Transactions and Proceedings of the New Zealand Institute 59: 232 - 286.","Stoliczka F. 1868. The Gastropoda. In: Blanford HF, ed. Cretaceous fauna of southern India, Vol. 2. Calcutta: Geological Survey of India, 1 - 497."]}

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

CREMNOCONCHUS GLOBULUS SP. NOV. (FIGS 2E, 4F, 5K, L, 8, 11H–L, 12H–L, 13A, B) Types: Holotype ZSI / WGRS /IR.INV-2310 (Fig. 11I, J); 2 paratypes ZSI / WGRS /IR.INV-2311, 2312 (Fig. 11K); Lesser Kadambi Falls, Chikmagalur Dist., Karnataka, India (13.24384°N, 75.17056°E). Etymology: Latin globulus, globular, in reference to shell shape. 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. 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. 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. 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. 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). 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. 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).

CREMNOCONCHUS GLOBULUS SP. NOV. (FIGS 2E, 4F, 5K, L, 8, 11H–L, 12H–L, 13A, B) Types: Holotype ZSI / WGRS /IR.INV-2310 (Fig. 11I, J); 2 paratypes ZSI / WGRS /IR.INV-2311, 2312 (Fig. 11K); Lesser Kadambi Falls, Chikmagalur Dist., Karnataka, India (13.24384°N, 75.17056°E). Etymology: Latin globulus, globular, in reference to shell shape. 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. 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. 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. 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. 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). 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. 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)., Published as part of 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), pp. 93-135 in Zoological Journal of the Linnean Society 167 (1) on page 121, DOI: 10.1111/j.1096-3642.2012.00875.x, http://zenodo.org/record/5292273

Copyright 2012 Shephard et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited., peer-reviewed, The life history characteristics of some elasmobranchs make them particularly vulnerable to fishing mortality; about a third of all species are listed by the IUCN as Threatened or Near Threatened. Marine Protected Areas (MPAs) have been suggested as a tool for conservation of elasmobranchs, but they are likely to be effective only if such populations respond to fishing impacts at spatial-scales corresponding to MPA size. Using the example of the Celtic Sea, we modelled elasmobranch biomass (kg h21) in fisheries-independent survey hauls as a function of environmental variables and ‘local’ (within 20 km radius) fishing effort (h y21) recorded from Vessel Monitoring Systems data. Model selection using AIC suggested strongest support for linear mixed effects models in which the variables (i) fishing effort, (ii) geographic location and (iii) demersal fish assemblage had approximately equal importance in explaining elasmobranch biomass. In the eastern Celtic Sea, sampling sites that occurred in the lowest 10% of the observed fishing effort range recorded 10 species of elasmobranch including the critically endangered Dipturus spp. The most intensely fished 10% of sites had only three elasmobranch species, with two IUCN listed as Least Concern. Our results suggest that stable spatial heterogeneity in fishing effort creates de facto refugia for elasmobranchs in the Celtic Sea. However, changes in the present fisheries management regime could impair the refuge effect by changing fisher’s behaviour and displacing effort into these areas., The research leading to these results has received funding from the European Community’s Seventh Framework Programme (FP7/2007–2013) under grant agreement MYFISH number 289257. David G. Reid also acknowledges funding from a Beaufort Marine Research Award, carried out under the Sea Change Strategy and the Strategy for Science Technology and Innovation (2006–2013), with the support of the Marine Institute, funded under the Marine Research Sub-Programme of the National Development Plan 2007–2013. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.

Subgenus Amerolittorina Reid, 2009 Echinolittorina (Amerolittorina) Reid, 2009: 33–35 (type species by original designation Trochus ziczac Gmelin, 1791). Diagnosis: Shell with spiral ribs and grooves, sometimes a pseudumbilicus, never granulose or nodulose; colour white with spots, oblique axial or spiral lines of brown to black; aperture brown with white band at base and usually also at shoulder. Penis usually bifurcate with mamilliform gland and penial glandular disc, but either or both may be absent. Pallial oviduct: copulatory bursa opens at posterior or anterior end of straight section. Atlantic and eastern Pacific. (Diagnosis from Reid 2009.) Remarks: This subgenus is phylogenetically distinct, but there is no diagnostic morphological synapomorphy (Reid 2009). It is restricted to the Atlantic and eastern Pacific Oceans, and consists of several morphologically and phylogenetically distinct complexes of species: the porcata group (6 species in the eastern Pacific); E. peruviana (Lamarck, 1822) (Peru and Chile); the punctata group (3 species in the eastern Atlantic); and the ziczac group (14 species in the eastern Pacific and Atlantic) (Reid 2009). Each group is monophyletic, but their relationships are not resolved in the most recent phylogenetic analysis (Williams & Duda 2008). The eastern Atlantic members are the three species in the punctata group, and the sister species of E. ziczac (Gmelin, 1791), described below. Since the early account by Adanson (1757) and its formal description by Gmelin (1791), E. punctata has been a well-known species in the Mediterranean and northwest Africa. Dunker (1845, 1853) introduced the name Litorina pulchella for colourful specimens from Angola, and some subsequent authors recognized this as distinct (Philippi 1847; Weinkauff 1882, 1883). Specimens from the Cape Verde Islands were usually assigned to one or other of these species, but Reeve (1857) considered them distinct (as L. guttata). From the late nineteenth century the punctata group has widely been considered to comprise a single species (Tryon 1887; Dautzenberg 1912; Nicklès 1950; Rosewater 1970, 1981), extending from the Mediterranean to Angola. Reid (2002) suggested on morphological grounds that three species were present and resurrected the name pulchella. This conclusion was subsequently confirmed by DNA sequence analysis (Williams & Reid 2004; Williams & Duda 2008). In addition to the similarity of their shells, members of the E. punctata group share an unusually small first loop of the albumen gland. Shells of the new species E. soroziczac from the islands of the Gulf of Guinea are superficially similar to those of the E. punctata group and were included with them until Reid (2002) pointed out their relationship with E. ziczac., Published as part of Reid, David G., 2011, 2974, pp. 1-65 in Zootaxa 2974 on pages 15-16, {"references":["Reid, D. G. (2009) The genus Echinolittorina Habe, 1956 (Gastropoda: Littorinidae) in the western Atlantic Ocean. Zootaxa, 2184, 1 - 103.","Gmelin, J. F. (1791) Caroli a Linne ... Systema Naturae per Regna Tria Naturae. 13 th ed. Vol. 1, part 6. Lipsiae.","Lamarck, J. B. P. A. de M. de (1822) Histoire Naturelle des Animaux sans Vertebres. Vol. 7. Paris.","Williams, S. T. & Duda, T. F. (2008) Did tectonic activity stimulate Oligo-Miocene speciation in the Indo-West Pacific? Evolution, 62, 1618 - 1634.","Adanson, M. (1757) Histoire Naturelle du Senegal. Coquillages. C. J. B. Bauche, Paris, 190 + 96 + 275 pp.","Dunker, W. (1845) Diagnoses molluscorum quorundam novorum, quae ex itinere ad oras Africae occidentalis reportavit cl. G. Tams, Med. Dr. Zeitschrift fur Malakozoologie, 2, 163 - 168.","Dunker, W. (1853) Index molluscorum, quae in itinere ad Guineam inferiorem collegit Georgius Tams Med. Dr. Novitates Conchologicae Suppl. 2, 1 - 74.","Reeve, L. A. (1857 - 1858) Monograph of the genus Littorina. In: Reeve, L. A. (Ed.), Conchologia Iconica. L. Reeve, London, Pls 1 - 16 (1857), 17 - 18 (1858).","Tryon, G. W. (1887) Manual of Conchology. Vol. 9. Philadelphia, 488 pp.","Dautzenberg, P. (1912) Mission Gruvel sur la cote occidentale d'Afrique (1909 - 1910): mollusques marins. Annales de l'Institut Oceanographique, 5 (3), 1 - 111.","Nickles, M. (1950) Mollusques testaces marins de la cote occidentale d'Afrique. Manuels Ouest-Africains, 2, 1 - 269.","Rosewater, J. (1970) The family Littorinidae in the Indo-Pacific. Part I. The subfamily Littorininae. Indo-Pacific Mollusca, 2, 417 - 506.","Rosewater, J. (1981) The family Littorinidae in tropical West Africa. Atlantide Report, 13, 7 - 48.","Williams, S. T. & Reid, D. G. (2004) Speciation and diversity on tropical rocky shores: a global phylogeny of snails of the genus Echinolittorina. Evolution, 58, 2227 - 2251."]}

Subgenus Echinolittorina Habe, 1956 Nodilittorina (Echinolittorina) Habe, 1956: 96–99 (type by original designation Litorina tuberculata Menke, 1828; as Echinolittiorina in error, p. 96). Diagnosis: Shell nodulose and spirally striate, occasionally a pseudumbilicus; one or two white lines in aperture. Penis bifurcate, with penial glandular disc and 1–2 mamilliform glands. Pallial oviduct: additional glandular material present in a swelling around egg groove at anterior end of straight section; copulatory bursa opens at posterior end of straight section. Tropical Atlantic and eastern Pacific. (Diagnosis modified from Reid 2009.) Remarks: Genetic data show that the three eastern Atlantic members of this subgenus, the E. granosa group (E. granosa, E. miliaris, E. helenae), form a clade (Williams & Reid 2004; Williams & Duda 2008; Reid 2009). There has been a history of taxonomic confusion between them and, together with E. vermeiji (Bandel & Kadolsky, 1982), all have at times been considered subspecies or synonyms of the first species to be named, E. miliaris. Their distributions are allopatric and, despite intraspecific variation, their shells are generally distinct. Anatomical differences among them are minimal; their penes are similar, but E. miliaris has a uniquely large lower chamber of the spiral part of the pallial oviduct. Tectarius striatus is another striate or granulose littorinid from the eastern Atlantic archipelagos, and has been confused with the members of the E. granosa group. Occasionally all the nodulose Echinolittorina species and T. striatus have been synonymized as Nodilittorina tuberculata (García-Talavera 1983), which is a Caribbean species. Like other nodulose members of the genus Echinolittorina (Reid, 2002b, 2007, 2009) these three species each show a remarkable range of intraspecific variation. The most extreme nodulose forms are usually smaller with a taller spire, whereas those with weak or absent nodules (including non-nodulose ‘striate’ forms in E. helenae) are larger and broader. The parallels with shell variation in other Echinolittorina species (especially E. australis (Gray, 1826); see review by Reid 2007) suggest an ecophenotypic basis and a likely correlation of nodulose sculpture with slow growth in open sunny conditions without strong wave splash. A similar range of variation is found in the distantly related Tectarius striatu s from the Cape Verde Islands, Canary Islands and Azores (Reid 1996), and shells of E. helenae can be almost indistinguishable from that species. The characters of all four are compared in Table 2. There is a superficial similarity between the nodulose shell forms of the eastern Atlantic E. (Echinolittorina) species and those of some members of E. (Granulilittorina) in the Indo-West Pacific. One useful feature for rapid discrimination between these groups is the presence of two pale bands in the aperture (the universal basal band and another at the shoulder) in many (but not all) shells of E. (Echinolittorina); the shoulder band is never present in the subgenus Granulilittorina., Published as part of Reid, David G., 2011, 2974, pp. 1-65 in Zootaxa 2974 on page 36, {"references":["Habe, T. (1956) Notes on the systematic position of three American sea shells. Venus, 19, 95 - 100.","Menke, K. T. (1828) Synopsis Methodica Molluscorum. H. Gelpke, Pyrmonti, xii + 91 pp.","Reid, D. G. (2009) The genus Echinolittorina Habe, 1956 (Gastropoda: Littorinidae) in the western Atlantic Ocean. Zootaxa, 2184, 1 - 103.","Williams, S. T. & Reid, D. G. (2004) Speciation and diversity on tropical rocky shores: a global phylogeny of snails of the genus Echinolittorina. Evolution, 58, 2227 - 2251.","Williams, S. T. & Duda, T. F. (2008) Did tectonic activity stimulate Oligo-Miocene speciation in the Indo-West Pacific? Evolution, 62, 1618 - 1634.","Bandel, K. & Kadolsky, D. (1982) Western Atlantic species of Nodilittorina (Gastropoda: Prosobranchia): comparative morphology and its functional, ecological, phylogenetic and taxonomic implications. Veliger, 25, 1 - 42.","Garcia-Talavera, F. (1983) Los moluscos gasteropodos anfiatlanticos (estudio paleo y biogeografico de las especies bentonicas litorales). Secretariado de Publicaciones de la Universidad de La Laguna Coleccion Monografias, 10, 1 - 351.","Reid, D. G. (2002 b) The genus Nodilittorina von Martens, 1897 (Gastropoda: Littorinidae) in the eastern Pacific Ocean, with a discussion of biogeographic provinces of the rocky-shore fauna. Veliger, 45, 85 - 169.","Reid, D. G. (2007) The genus Echinolittorina Habe, 1956 (Gastropoda: Littorinidae) in the Indo-West Pacific Ocean. Zootaxa, 1420, 1 - 161.","Gray, J. E. (1826) Mollusca. In: P. P. King (Ed.) Narrative of a Survey of the Intertropical and Western Coasts of Australia. Performed between the years 1818 and 1822. John Murray, London, pp. 474 - 496.","Reid, D. G. (1996) Systematics and Evolution of Littorina. Ray Society, London. x + 463 pp."]}

Genus Echinolittorina Habe, 1956 Nodilittorina (Echinolittorina) Habe, 1956: 96–99 (type species by original designation Litorina tuberculata Menke, 1828; cited as Echinolittiorina in error, p. 96). Diagnosis: Shell aragonitic; nodulose, granulose, spirally striate or entirely smooth; if smooth or striate then usually with dark axial stripes or axially aligned series of dashes; eroded parietal area may be present, sometimes a small pseudumbilicus; aperture dark with pale spiral band at base (i.e. anterior end) and sometimes another posteriorly. Cephalic tentacles with 2–3 longitudinal black lines, all black, or with broad transverse band at base distal to eye. Operculum paucispiral. Penis with blade-shaped filament (rarely swollen); base usually bifurcate (not so if mamilliform gland absent); usually a single mamilliform gland (sometimes absent, rarely 2); glandular disc of subepithelial glandular tissue usually present; penial vas deferens an open groove or rarely a closed tube. Rod-pieces of paraspermatozoa usually long and straight (rarely small and irregular, or curved). In pallial oviduct egg groove makes a simple loop through albumen gland, straight path through capsule gland and jelly gland; copulatory bursa opens in anterior or posterior position within straight section of pallial oviduct. Spawn a biconvex pelagic capsule with cupola-shaped upper side sculptured by concentric rings, containing single egg; development planktotrophic. Outer marginal radular tooth with flange at inside and usually also at outside of base. Distribution worldwide tropical and warm temperate. (Diagnosis from Reid 2009.) Remarks: For taxonomic history, see Reid (2007). Monographs are available of the eastern Pacific species [Reid 2002a; as ‘ Nodilittorina ’, but Austrolittorina fernandezensis (Rosewater, 1970) and A. araucana (d’Orbigny, 1840) are now excluded (Williams et al. 2003)], Indo-Pacific species (Reid 2007) and western Atlantic species (Reid 2009).

Subgenus Fossarilittorina Rosewater, 1981 Littorina (Fossarilittorina) Rosewater, 1981: 29 (type species by original designation Phasianella meleagris [Beck in] Potiez & Michaud, 1838). Diagnosis: Shell smooth, with microstriae and a single incised line at periphery; pseudumbilicus adjacent to columella. Penis simple, lacking mamilliform glands and glandular disc, sperm duct closed as a tube. Pallial oviduct contains an additional gland around egg groove at posterior end of straight section (anterior to translucent capsule gland); copulatory bursa opens at anterior end of straight section. Radula lacks flange on outer side of base of outer marginal tooth. Caribbean Sea, Gulf of Mexico, Gulf of Guinea to Angola. (Diagnosis from Reid 2009.) Remarks: For taxonomic history, see Reid (2009). This subgenus is restricted to the Atlantic Ocean, with two species in the western Atlantic and one, described below, in West Africa. The small size, umbilicate shell and relatively low-shore (mid- to low-eulittoral) habitat are unusual in the genus Echinolittorina., Published as part of Reid, David G., 2011, 2974, pp. 1-65 in Zootaxa 2974 on page 10, {"references":["Rosewater, J. (1981) The family Littorinidae in tropical West Africa. Atlantide Report, 13, 7 - 48.","Potiez, V. L. V. & Michaud, A. L. G. (1838) Galerie des Mollusques, ou Catalogue Methodique, Descriptif et Raisonne des Mollusques et Coquilles du Museum de Douai. Vol. 1. J. - B. Bailliere, Paris, xxxi + 560 pp.","Reid, D. G. (2009) The genus Echinolittorina Habe, 1956 (Gastropoda: Littorinidae) in the western Atlantic Ocean. Zootaxa, 2184, 1 - 103."]}

Genus Echinolittorina Habe, 1956 Nodilittorina (Echinolittorina) Habe, 1956: 96–99 (type species by original designation Litorina tuberculata Menke, 1828; cited as Echinolittiorina in error, p. 96). Diagnosis: Shell aragonitic; nodulose, granulose, spirally striate or entirely smooth; if smooth or striate then usually with dark axial stripes or axially aligned series of dashes; eroded parietal area may be present, sometimes a small pseudumbilicus; aperture dark with pale spiral band at base (i.e. anterior end) and sometimes another posteriorly. Cephalic tentacles with 2–3 longitudinal black lines, all black, or with broad transverse band at base distal to eye. Operculum paucispiral. Penis with blade-shaped filament (rarely swollen); base usually bifurcate (not so if mamilliform gland absent); usually a single mamilliform gland (sometimes absent, rarely 2); glandular disc of subepithelial glandular tissue usually present; penial vas deferens an open groove or rarely a closed tube. Rod-pieces of paraspermatozoa usually long and straight (rarely small and irregular, or curved). In pallial oviduct egg groove makes a simple loop through albumen gland, straight path through capsule gland and jelly gland; copulatory bursa opens in anterior or posterior position within straight section of pallial oviduct. Spawn a biconvex pelagic capsule with cupola-shaped upper side sculptured by concentric rings, containing single egg; development planktotrophic. Outer marginal radular tooth with flange at inside and usually also at outside of base. Distribution worldwide tropical and warm temperate. (Diagnosis from Reid 2009.) Remarks: For taxonomic history, see Reid (2007). Monographs are available of the eastern Pacific species [Reid 2002a; as ‘ Nodilittorina ’, but Austrolittorina fernandezensis (Rosewater, 1970) and A. araucana (d’Orbigny, 1840) are now excluded (Williams et al. 2003)], Indo-Pacific species (Reid 2007) and western Atlantic species (Reid 2009)., Published as part of Reid, David G., 2011, 2974, pp. 1-65 in Zootaxa 2974 on page 10, {"references":["Habe, T. (1956) Notes on the systematic position of three American sea shells. Venus, 19, 95 - 100.","Menke, K. T. (1828) Synopsis Methodica Molluscorum. H. Gelpke, Pyrmonti, xii + 91 pp.","Reid, D. G. (2009) The genus Echinolittorina Habe, 1956 (Gastropoda: Littorinidae) in the western Atlantic Ocean. Zootaxa, 2184, 1 - 103.","Reid, D. G. (2007) The genus Echinolittorina Habe, 1956 (Gastropoda: Littorinidae) in the Indo-West Pacific Ocean. Zootaxa, 1420, 1 - 161.","Reid, D. G. (2002 a) Morphological review and phylogenetic analysis of Nodilittorina Gastropoda: Littorinidae). Journal of Molluscan Studies, 68, 259 - 281.","Rosewater, J. (1970) The family Littorinidae in the Indo-Pacific. Part I. The subfamily Littorininae. Indo-Pacific Mollusca, 2, 417 - 506.","Williams, S. T., Reid, D. G. & Littlewood, D. T. J. (2003) A molecular phylogeny of the Littorininae (Gastropoda: Littorinidae): unequal evolutionary rates, morphological parallelism and biogeography of the Southern Ocean. Molecular Phylogenetics and Evolution, 28, 60 - 86."]}

Subgenus Echinolittorina Habe, 1956 Nodilittorina (Echinolittorina) Habe, 1956: 96–99 (type by original designation Litorina tuberculata Menke, 1828; as Echinolittiorina in error, p. 96). Diagnosis: Shell nodulose and spirally striate, occasionally a pseudumbilicus; one or two white lines in aperture. Penis bifurcate, with penial glandular disc and 1–2 mamilliform glands. Pallial oviduct: additional glandular material present in a swelling around egg groove at anterior end of straight section; copulatory bursa opens at posterior end of straight section. Tropical Atlantic and eastern Pacific. (Diagnosis modified from Reid 2009.) Remarks: Genetic data show that the three eastern Atlantic members of this subgenus, the E. granosa group (E. granosa, E. miliaris, E. helenae), form a clade (Williams & Reid 2004; Williams & Duda 2008; Reid 2009). There has been a history of taxonomic confusion between them and, together with E. vermeiji (Bandel & Kadolsky, 1982), all have at times been considered subspecies or synonyms of the first species to be named, E. miliaris. Their distributions are allopatric and, despite intraspecific variation, their shells are generally distinct. Anatomical differences among them are minimal; their penes are similar, but E. miliaris has a uniquely large lower chamber of the spiral part of the pallial oviduct. Tectarius striatus is another striate or granulose littorinid from the eastern Atlantic archipelagos, and has been confused with the members of the E. granosa group. Occasionally all the nodulose Echinolittorina species and T. striatus have been synonymized as Nodilittorina tuberculata (García-Talavera 1983), which is a Caribbean species. Like other nodulose members of the genus Echinolittorina (Reid, 2002b, 2007, 2009) these three species each show a remarkable range of intraspecific variation. The most extreme nodulose forms are usually smaller with a taller spire, whereas those with weak or absent nodules (including non-nodulose ‘striate’ forms in E. helenae) are larger and broader. The parallels with shell variation in other Echinolittorina species (especially E. australis (Gray, 1826); see review by Reid 2007) suggest an ecophenotypic basis and a likely correlation of nodulose sculpture with slow growth in open sunny conditions without strong wave splash. A similar range of variation is found in the distantly related Tectarius striatu s from the Cape Verde Islands, Canary Islands and Azores (Reid 1996), and shells of E. helenae can be almost indistinguishable from that species. The characters of all four are compared in Table 2. There is a superficial similarity between the nodulose shell forms of the eastern Atlantic E. (Echinolittorina) species and those of some members of E. (Granulilittorina) in the Indo-West Pacific. One useful feature for rapid discrimination between these groups is the presence of two pale bands in the aperture (the universal basal band and another at the shoulder) in many (but not all) shells of E. (Echinolittorina); the shoulder band is never present in the subgenus Granulilittorina.

Mechanical treatment of chemicals by simply grinding them together in a ball mill has been identified as a clean and environmentally-friendly synthetic methodology that meets the requirements of green chemistry (1). As such, mechanochemistry has recently opened new opportunities for the design and construction of new materials, e.g. coordination polymers, pharmaceutical derivatives and porous metal-organic frameworks (MOFs) (2). We envisage that, in combination with modern solid-state analytical methods, automated ball-milling can provide an almost entirely solvent-free route from the synthesis and analysis to the full structural characterisation of a product. In this contribution we present a mechanochemical approach for the selective synthesis of simple metal-organic frameworks. Solid-state methodology based on the concept of liquid-assisted grinding (LAG) (3) allows a rapid screening for different stoichiometric compositions of MOFs with cyanoguanidine as a heteroditopic ligand (4). The products obtained by mechanosynthesis have been further characterised by solid-state analytical tools such as FTIR-ATR, solid-state NMR and structure solution from X-ray powder diffraction data, eliminating the need for bulk solvents in our research laboratory.

Mechanical treatment of chemicals by simply grinding them together in a ball mill has been identified as a clean and environmentally-friendly synthetic methodology that meets the requirements of green chemistry (1). As such, mechanochemistry has recently opened new opportunities for the design and construction of new materials, e.g. coordination polymers, pharmaceutical derivatives and porous metal-organic frameworks (MOFs) (2). We envisage that, in combination with modern solid-state analytical methods, automated ball-milling can provide an almost entirely solvent-free route from the synthesis and analysis to the full structural characterisation of a product. In this contribution we present a mechanochemical approach for the selective synthesis of simple metal-organic frameworks. Solid-state methodology based on the concept of liquid-assisted grinding (LAG) (3) allows a rapid screening for different stoichiometric compositions of MOFs with cyanoguanidine as a heteroditopic ligand (4). The products obtained by mechanosynthesis have been further characterised by solid-state analytical tools such as FTIR-ATR, solid-state NMR and structure solution from X-ray powder diffraction data, eliminating the need for bulk solvents in our research laboratory.

We are witnessing the problems of insufficient energy resources and environment pollution that have imposed a great challenge upon modern chemistry science. In a search for methodologies that would enable a clean and environmentally-friendly, while still economically accessible approach to design and construction of new materials, mechanochemistry is recognised as a method of choice. Recent discoveries in the field of solid-state reactivity such as liquid-assisted grinding (LAG)have promoted mechanosynthesis as genuinely a „green chemistry“ method far superior to the classical solution-based techniques. Moreover, an integration of mechanochemistry with modern solid-state analytical tools (structure solution from X-ray powder diffraction data - PXRD and solid-state NMR) [3] allows the entire search from synthesis and analysis to structural characterisation of a product to be conducted under a fully solvent-free conditions. In this presentation, a mechanochemical approach for the synthesis of zinc and cadmium complexes of cyanoguanidine will be described. We demonstrate the power of mechanosynthesis to screen for different stoichiometric compositions of metal-organic polymers and the use of a heteroditopic ligand to control the imensionality of such materials. By coupling the grinding synthetic methodology with PXRD and solid-state NMR characterisation tools we have achieved a complete implementation of the solvent-free paradigm in laboratory research.

Genus Echinolittorina Habe, 1956 Nodilittorina (Echinolittorina) Habe, 1956: 96–99 (type by original designation Litorina tuberculata Menke, 1828; cited as Echinolittiorina in error, p. 96). Diagnosis: Shell aragonitic; nodulose, granulose, spirally striate or entirely smooth; if smooth or striate then usually with dark axial stripes or axially aligned series of dashes; eroded parietal area may be present, sometimes a small pseudumbilicus; aperture dark with pale spiral band at base (i.e. anterior end) and sometimes another posteriorly. Cephalic tentacles with 2–3 longitudinal black lines, all black, or with broad transverse band at base distal to eye. Operculum paucispiral. Penis with blade-shaped filament (rarely swollen); base usually bifurcate (not so if mamilliform gland absent); usually a single mamilliform gland (sometimes absent, rarely 2); glandular disc of subepithelial glandular tissue usually present; penial vas deferens an open groove or rarely a closed tube. Rod-pieces of paraspermatozoa usually long and straight (rarely small and irregular, or curved). In pallial oviduct egg groove makes a simple loop through albumen gland, straight path through capsule gland and jelly gland; copulatory bursa opens in anterior or posterior position within straight section of pallial oviduct. Spawn a biconvex pelagic capsule with cupola-shaped upper side sculptured by concentric rings, containing single egg; development planktotrophic. Outer marginal radular tooth with flange at inside and usually also at outside of base. Distribution worldwide tropical and warm temperate. (Modified from Reid 2007.) Remarks: The taxonomic history of this genus has been summarized by Reid (2007). Since the molecular phylogeny of Echinolittorina species by Williams & Reid (2004) a revised analysis with addition of new sequences (including the new species described herein, making a complete analysis of all 60 known members worldwide) has been published (Williams & Duda 2008; Fig. 1). Monographs are available of the eastern Pacific species [Reid 2002b; as ‘ Nodilittorina ’, but Austrolittorina fernandezensis (Rosewater, 1970) and A. araucana (d’Orbigny, 1840) are now excluded (Williams et al. 2003)] and of the Indo-Pacific species (Reid 2007)., Published as part of Reid, David G., 2009, The genus Echinolittorina Habe, 1956 (Gastropoda: Littorinidae) in the western Atlantic Ocean 2184, pp. 1-103 in Zootaxa 2184 (1) on page 14, DOI: 10.11646/zootaxa.2184.1.1, http://zenodo.org/record/5318099, {"references":["Habe, T. (1956) Notes on the systematic position of three American sea shells. Venus, 19, 95 - 100.","Menke, K. T. (1828) Synopsis Methodica Molluscorum. H. Gelpke, Pyrmonti, xii + 91 pp.","Reid, D. G. (2007) The genus Echinolittorina Habe, 1956 (Gastropoda: Littorinidae) in the Indo-West Pacific Ocean. Zootaxa, 1420, 1 - 161.","Williams, S. T. & Reid, D. G. (2004) Speciation and diversity on tropical rocky shores: a global phylogeny of snails of the genus Echinolittorina. Evolution, 58, 2227 - 2251.","Williams, S. T. & Duda, T. F. (2008) Did tectonic activity stimulate Oligo-Miocene speciation in the Indo-West Pacific? Evolution, 62, 1618 - 1634.","Reid, D. G. (2002 b) The genus Nodilittorina von Martens, 1897 (Gastropoda: Littorinidae) in the eastern Pacific Ocean, with a discussion of biogeographic provinces of the rocky-shore fauna. Veliger, 45, 85 - 169.","Rosewater, J. (1970) The family Littorinidae in the Indo-Pacific. Part I. The subfamily Littorininae. Indo-Pacific Mollusca, 2, 417 - 506.","Williams, S. T., Reid, D. G. & Littlewood, D. T. J. (2003) A molecular phylogeny of the Littorininae (Gastropoda: Littorinidae): unequal evolutionary rates, morphological parallelism and biogeography of the Southern Ocean. Molecular Phylogenetics and Evolution, 28, 60 - 86."]}