The radula of the Late Cretaceous scaphitid ammonite Rhaeboceras halli (Meek and Hayden, 1856)

Abstract: Radular teeth occur between the jaws in two specimens of the Late Cretaceous scaphitid ammonite Rhaeboceras halli (Meek and Hayden, 1856) from the Western Interior of the United States. The detailed morphology of the teeth has been revealed by propagation phase contrast X‐ray synchrotron microtomography. Each row of the radula of R. halli consists of a total of seven teeth (a central rachidian, two pairs of lateral and one pair of marginal teeth), as in other known ammonoid radulae, although the central tooth could not be confirmed in the specimens examined. The lateral teeth are multicuspid and robust, and the marginal teeth are long (4.6 mm) and slender. In overall morphology, the heterodont and ctenoglossan radula of R. halli is similar that of Jurassic and Cretaceous ammonites with the same aptychus‐type lower jaw, that is, the Aptychophora. This discovery reveals the range of variation in radular morphology, which could be related to ecological or phylogenetic factors. It also invalidates the hypothesis that the hook‐like structures in R. halli previously described are radular elements.     

Fine Structure of the Accessory Boring Organ of the Gastropod, Urosalpinx

Penetration of shell by the muricid gastropod, Urosalpinx cinereafollyensis, is accomplished by successive alternating periodsof radular rasping and chemical activity by the accessory boringorgan (ABO) . This paper reports on thehistology of the ventralsecretory disc of the ABO in relation to the functional stateof the organ. The secretory disc is composed of tall columnarepithelial cells which are arranged in groups and only forma continuum over the ABO surface at their distalmost end. Theinterstitial spaces between the epithelial cell groups containmuscles and capillaries and also form the link between the distallyopen capillaries and the sinus of the ABO stalk. Abundant nervefibers are found interstitially as well as in close relationshipto the epithelium. The epithelial cells possess cytological features characteristicof secretory cells. The increased differentiation especiallyof the Golgi complex during periods of active boring indicatesthat the cells elaborate a product which plays a rolein thechemical dissolution of the shell of prey. The cells also appearcapable of regulating secretory overproduction and of storingglycogen during inactive periods for subsequent use.     

Two new trochids of the genus <Emphasis Type="Italic">Antimargarita</Emphasis> (Gastropoda: Vetigastropoda: Trochidae) from the Bellingshausen Sea and South Shetland Islands,Antarctica

Two new trochids of the genus Antimargarita, A. powelli and A. bentarti, from the Antarctic waters are described here. A. powelli, from the Bellingshausen Sea, is distinguished by its rounded whorls, numerous spiral cords, a radula with seven lateral teeth at each side of the rachidian, and an epipodium with eight pairs of tentacles. A. bentarti, from the South Shetland Islands, is characterized by having a shell outline gradated by prominent primary spiral cords, a radula with five lateral teeth at each side of the rachidian and an epipodium with six tentacles on the left side. The diagnostic features for Antimargarita are redefined considering both shell and anatomical features and its suprageneric placement is discussed.     

RADULAR PRODUCTION RATES IN TWO SPECIES OF LACUNA TURTON (GASTROPODA: LITTORINIDAE)

The molluscan radula is a dynamic organ, both in terms of itsuse and production. New rows of teeth are constantly producedat the posterior end of the radula, while older, worn teethare shed anteriorly, producing a dynamic equilibrium. We useda cold-shock to mark the radular ribbon and measure tooth rowproduction rates in two gastropod species, Lacuna vincta (Montagu)and L. vanegata Carpenter. We found that the average tooth rowproduction rate at 10–11°C did not differ betweenthese two species, and was 2.94 (SE = 0.002) rows per day forLacuna vincta and 2.97 (SE = 0 002) for L. vanegata Inter-individualvariability in production rate was very low, and was correlatedwith shell length, smaller individuals had slightly higher productionrates. The total length of the radular nbbon varied greatlyamong individuals, ranging from 47 to 94 (2.57 to 5.68 mm) rowsin L vincta and 53 to 99 rows (2.80 to 7.14 mm) in L vanegata,and was only somewhat correlated with the length of the shelLThis great variability will result in large differences amongindividuals in the time it takes to replace the radula totally,from 14.96 to 35.44 days in L vincta and from 17 43 to 39 69days in L. vanegata. (Received 1 September 1995; accepted 20 November 1995)     

两种石磺齿舌形态差异分析

显微观察了瘤背石磺(Onchidiumstruma)和石磺(O. verruculatum)齿舌的形态结构。运用差异系数法对两种石磺齿舌参数进行比较分析。利用SPSS10.0对瘤背石磺、石磺齿舌参数(齿舌长、齿舌头宽、齿舌中宽、齿舌尾宽、横列数、每排最少齿片数和每排最多齿片数)与个体参数(体长、体宽、体高、足长、足宽和体重)作回归分析。结果表明,两种石磺齿舌都很发达,外观呈长统靴状;齿片排成许多横列,每一横列均有中央齿一枚,侧齿若干无缘齿;两种石磺的齿舌头宽、齿舌中宽和齿舌尾宽差异极显著,但差异系数小于1.28,认为两种石磺的齿片形态存在明显的种间差异,但齿舌参数不适合作为石磺属贝类的分类依据;瘤背石磺的体宽和石磺的体重在评估各自齿舌生物学性状方面起到比较重要的作用。     

RADULAR VARIATION IN TWO SPECIES OF SPONGE-RASPING DORID NUDIBRANCHS

The damp live weight of specimens of Archidoris montereyensisand Anisodoris nobilis was found to be positively correlated( = 0.05) to the number of teeth per row, the number of rowsin the radula and the length of teeth. Covariance analyses ofthe regressions of the first two radular characteristics toweight failed to statistically separate the two species. Theseresults argue against the utility of radular information astaxonomic characters in sponge-rasping dorids. The increase in tooth size with increasing animal size was foundto be statistically divergent for these two species and wasinterpreted as being consistent with the feeding biology ofthese two species. (Received 10 March 1977;     

Indirect evidence for ecophenotypic plasticity in radular dentition of Littoraria species (Gastropoda: Littorinidae)

In examination of radulae from all but one of the 36 speciesof the littorinid genus Littoraria we found extraordinary intraspecificvariation in those occurring on a range of substrates. Radulaefrom rock showed a less well developed `hood' on the rachidiantooth, a strikingly enlarged major cusp on each of the fivecentral teeth, fewer cusps on the outer marginal teeth and theradular ribbon was longer, when compared with radulae of conspecificsfrom plant substrates. The radulae of species found exclusivelyon rock differed in similar ways from those restricted to plantsubstrates (mangroves, driftwood and saltmarsh). We suggestthat this may be an example of phenotypic plasticity of radularform, induced by substrate and/or diet, as recently shown experimentallyin another littorinid genus. The mechanism of inducible plasticitydeserves further study. Ecotypic variation in the radula maybe widespread in littorinids, and radular characters shouldtherefore be used with caution in studies of taxonomy, phylogenyand adaptation. (Received 20 July 1998; accepted 10 November 1998)     

Seasonal aberrant radular formation in Thais bronni (Dunker) and T. clavigera (Küster) (Gastropoda : Muricidae)

Radulae of Thais bronni (Dunker) and T. clavigera (Küster) were examined at Mukaishima Island for a period of 2 yr, 1982 to 1984. Radulae of both species are similar in morphology, both having the basic pentacuspid rachidian plan. Sexual dimorphism of the radula was not observed, but rachidian tooth changes morphologically in different growth stages. Seasonal conditions affect the size and shape of the radula; in winter it is clearly malformed and strikingly thin. These aberrant parts of the radula comprised some dozens of rows, in which only several extremely thin rows exist. Results of experiments using T. clavigera under different water temperature conditions showed that the radula is rarely produced below 10 °C and that rate of radular production and replacement increases with increase in temperature. These results suggest that in the field the radula of these species is replaced entirely 2–2.5 times per year and 10–15 times during the life of the animal.     

Radula formation in two species of Conoidea (Gastropoda)

The radula is the basic feeding structure in gastropod molluscs and exhibits great morphological diversity that reflects the exceptional anatomical and ecological diversity occurring in these animals. This uniquely molluscan structure is formed in the blind end of the radular sac by specialized cells (membranoblasts and odontoblasts). Secretion type, and the number and shape of the odontoblasts that form each tooth characterize the mode of radula formation. These characteristics vary in different groups of gastropods. Elucidation of this diversity is key to identifying the main patterns of radula formation in Gastropoda. Of particular interest would be a phylogenetically closely related group that is characterized by high variability of the radula. One such group is the large monophyletic superfamily Conoidea, the radula of which is highly variable and may consist of the radular membrane with five teeth per row, or the radular membrane with only two or three teeth per row, or even just two harpoon-like teeth per row without a radular membrane. We studied the radulae of two species of Conoidea (Clavus maestratii Kilburn, Fedosov & Kantor, 2014 [Drilliidae] and, Lophiotoma acuta (Perry, 1811) [Turridae]) using light and electron microscopy. Based on these data and previous studies, we identify the general patterns of the radula formation for all Conoidea: the dorsolateral position of two groups of odontoblasts, uniform size, and shape of odontoblasts, folding of the radula in the radular sac regardless of the radula configuration. The morphology of the subradular epithelium is most likely adaptive to the radula type.     

Age determination and estimation of larval period in field caught abalone (Haliotis discus hannai Ino 1953) larvae and newly metamorphosed post-larvae by counts of radular teeth rows

We developed an age determination method for larval and newly metamorphosed post-larval abalone Haliotis discus hannai in a laboratory experiment and determined the age of field caught individuals. Laboratory experiments showed that competent veliger larvae (4 days after fertilization) had a radula and regularly added rows of radular teeth with age in the absence of metamorphosis. Under environmentally relevant temperatures (17-22 °C), the number of rows of radular teeth increased linearly with age, but slopes of the regression lines were different among temperatures. Rows of radular teeth were added more slowly at lower temperatures. The effect of temperature on the development rate of the radula was quantified by the regression and the temperature coefficient, Q₁₀. The radular development of newly metamorphosed post-larvae, which had not acquired a peristomal shell (adult shell), was comparable with that of veliger larvae, although older post-larvae had a larger number of rows of radula than those of the same age of veliger larvae. From these results, an age determination method of veliger larvae and newly metamorphosed post-larvae was established, using the number of rows of radular teeth. The age of veliger larvae and newly metamorphosed post-larvae was determined by the age determination method for samples collected in August to October of 2003 and 2004 for which the thermal history of the coastal water of Miyagi Prefecture Japan was available. Only 9.1% of veliger larvae (n = 8) captured in the field had formed a radula and these were estimated to be 4-6 days old. The remaining 90.9% of larvae (n = 80) that had not formed a radula were classified as younger than 4 days old. All newly metamorphosed post-larvae (n = 24) that had metamorphosed on substrata were estimated to be 4-6 days old. Results of the field study indicate that these abalone metamorphosed within a few days after the acquisition of competence (4 days after fertilization) at this site, which has suitable crustose algal habitat.     

Some Aspects of Hole-Boring Predation by Octopus vulgaris

Octopus vulgaris prey upon many gastropod species by boringholes in the shell, weakening the prey with a venom, removingthe entire prey, and eating it. When offered Strombus raninusthe Octopus quickly grasped the conch with one or a few arms,checked for occupancy by inserting an arm tip into the aperture,and passed the shell under the web to the mouth. The shell washeld against the buccal mass by the circumoral suckers and raspedrepeatedly with the radula, repositioned, and rasped again.There were brief pauses of apparent inactivity between the periodsof active rasping. The shell was penetrated at an approximatemaximal rate of 1.25 mm per hour. The boreholes averaged 0.93mm in outer diameter, 0.47 mm in inner diameter and 0.88 mmin depth. The boreholes were extremely variable in shape, size,and position on the spire. There was a marked preference forindividual animals to bore in a particular sector of the spire.Apparently the animals orient the shells by using the lip asa point of reference because lipless shells had the boreholesrandomly distributed around the shell.     

The influence of season and the tidal cycle on division of labour by the radula during feeding in the estuarine brooding gastropod <Emphasis Type="Italic">Crepipatella dilatata</Emphasis> (Calyptraeidae)

The brooding gastropod Crepipatella dilatata can feed by scraping the substrate with the radula and by suspension-feeding, which also requires use of the radula. There is a “division of labour” for the radula among three discrete tasks associated with feeding: (1) removing mucous balls from the food pouch; (2) transferring the mucous cord from the neck channel to the mouth (both components of suspension-feeding); (3) scraping the substrate. We hypothesised that the proportion of time used for each feeding activity varies according to environmental conditions. Total radular activity in females was greatest at high tide and in summer. The rate of radular extrusion for ingesting the mucous cord varied seasonally and between brooding and non-brooding females. Non-brooding females exhibited higher rates of radular extrusion for ingesting the mucous cord and for scraping the substrate than did brooders. In females, radular activity in removing the mucous ball from the food pouch was strongly influenced by the tidal cycle during winter, reaching minimum values at low tide. Differences were recorded in substrate scraping among seasons and within tidal cycles, and among males, brooding females and non-brooding females. Brooding females displayed less rasping than non-brooders, since the area available for grazing was restricted by the egg mass. Throughout the year, including low salinity periods, males allocated a greater proportion of total radular activity to rasping than to removing the mucous ball or ingesting the mucous cord. The feeding behaviour of both males and females is modulated by salinity, but the principal determinants of radular activity are the mode of reproduction (brooding in females) and, in males, motility.     

COUPLING MOTOGRAPHIC AND SONOGRAPHIC RECORDING TO ASSESS FORAGING BEHAVIOUR OF PATELLA VULGATA

Motographic recording of locomotor activity and sonographicrecording of rasping were simultaneously obtained in the fieldfrom a group of Patella vulgata L. inhabiting a sheltered shore(Menai Bridge, North Wales). Each limpet was equipped with apiezoelectric transducer, a small amplifier, and a micro-lighterencased in dental acrylic and glued to the shell. A remote filter-amplifier-recorderdevice continuously recorded rasping noise through the activityperiods (nocturnal low tide), when individual movements werealso recorded using an automatic camera. Movements of experimentallimpets did not differ from those of specimens equipped withthe micro-lighter only. Locomotor activity was definitely concentratedduring the first (outgoing) and last (homing) parts of eachactivity phase. Rasping started when limpets remained exposedto air at ebb tide, 1–2 hours before leaving home, andoccurred during the whole period spent away from home. Raspingwas fairly variable in rate, consisting in non-rhythmic boutsseparated by recovery phases. No significant correlation wasobserved between rasping rate and speed variation during differentparts of each excursion. However, the number of rasps performedin each spatial unit of the path was strongly correlated withthe time spent by the limpet in that unit. The data suggestthat in Patella vulgata the spatial organisation of foragingwithin each excursion is based on changes in travel speed relatedto local food density but not on modulation of rasping ratein time. (Received 4 July 1993; accepted 2 August 1993)     

A new Georissa (Gastropoda: Neritopsina: Hydrocenidae) from a limestone cave in Malaysian Borneo

In a previous paper, we have reported on a new, troglobiticspecies of Georissa and its possible parapatric origin fromGeorissa saulae Benthem-Jutting, 1966, which occurs outsidethe cave system of Batu Sanaron, a limestone outcrop in Sabah,Malaysian Borneo. These analyses were based on genetic and morphometricdata. Here, we formally describe the new species adding anatomicaldata derived from dissections and histological serial sectionsas well as fine structural data of shell, operculum and radula,and compare it with its stem species. The two species differin shell and radular morphology as well as genital characters.Since we found anatomical differences between very closely relatedspecies, we assume that dissections would be of general usefor the taxonomy of Georissa and the remaining nominal genus-grouptaxa of the poorly known Hydrocenidae.     

DESCRIPTION OF EUBRANCHUS LINENSIS NEW SPECIES (NUDIBRANCHIA), WITH REMARKS ON DIAULY IN NUDIBRANCHS

A new species of nudibranch, Eurbranchus liensis, is described,from specimens collected from the southern Iberian Peninsula.The coloration of this species is characterized by the presenceof blood-red patches over the whole body and opaque white pigmentationon the tips of the cerata, rhinophores and oral tentacles, aswell as along the side of the foot. The jaws are denticulateand the triseriate radula has lateral teeth with a small secondarycusp. The rachidian tooth usually has four denticles on eachside. The reproductive system has a prostate and there is astylet at the end of the penis. The diaulic condition in theEubranchidae and in other families of nudibranchs is discussed. (Received 15 August 1989; accepted 24 February 1989)     

Main patterns of radula formation and ontogeny in Gastropoda

Gastropoda is morphologically highly variable and broadly distributed group of mollusks. Due to the high morphological and functional diversity of the feeding apparatus gastropods follow a broad range of feeding strategies: from detritivory to highly specialized predation. The feeding apparatus includes the buccal armaments: jaw(s) and radula. The radula comprises a chitinous ribbon with teeth arranged in transverse and longitudinal rows. A unique characteristic of the radula is its continuous renewal during the entire life of a mollusk. The teeth and the membrane are continuously synthesized in the blind end of the radular sac and are shifted forward to the working zone, while the teeth harden and are mineralized on the way. Despite the similarity of the general mechanism of the radula formation in gastropods, some phylogenetically determined features can be identified in different phylogenetic lineages. These mainly concern shape, size, and number of the odontoblasts forming a single tooth. The radular morphology depends on the shape of the formation zone and the morphology of the subradular epithelium. The radula first appears at the pre- and posttorsional veliger stages as an invagination of the buccal epithelium of the larval anterior gut. The larval radular sac is lined with uniform undifferentiated cells. Each major phylogenetic lineage is characterized by a specific larval radula type. Thus, the docoglossan radula of Patellogastropoda is characterized by initially three and then five teeth in a transverse row. The larval rhipidoglossan radula has seven teeth in a row with differentiation into central, lateral, and marginal teeth and later is transformed into the adult radula morphology by the addition of lateral and especially marginal teeth. The taenioglossan radula of Caenogastropoda is nearly immediately formed in adult configuration with seven teeth in a row.     

COMPARATIVE MORPHOLOGY OF RADULAR TEETH IN CONUS: OBSERVATIONS WITH SCANNING ELECTRON MICROSCOPY

Radular teeth of 22 Indo-Pacific species of the genus Conus(Neogastropoda: Toxoglossa) were compared. On morphologicalfeatures all can be related to one of three known feeding modes:piscivorous, vermivorous and molluscivorous. Observations arereported on the radular teeth of six piscivores, thirteen vermivoresand three molluscivores. The radular teeth of piscivores areof two general types. In the first, two barbs and a posteriorly-directedprocess with a recurved tip are found at the anterior end. Inthe second, two barbs are located at the anterior end and theshaft is serrated for most of its length. An enlarged posteriorregion (terminal knob) is present in the first and absent inthe second. Molluscivores possess radular teeth with two anteriorbarbs and in some species a serrated shaft or terminal knob.The radular teeth of vermivores, which show much greater interspecificvariation than those of piscivores or molluscivores, are characterizedby one or two anterior barbs and in most species a serratedregion near the apex. A forwardly-projecting cone (basal spur)is usually located on the terminal knob. Piscivores and molluscivoreslack such basal spurs. The radular teeth of Conus are used toconvey a potent venom and hold prey firmly during feeding. Previouslyundescribed morphological features are noted on the teeth ofC. obscurus and C. lividus. Figured here for the first timeare the radular teeth of C. abbreviatus, C. aureus, C. catus,C. litoglyphus, C. pennaceus, C. rattus and C. sponsalis. ^*Present address: Department of Paleontology, University ofCalifornia, Berkeley, California 94720, U.S.A. (Received 2 April 1979;     

FEEDING PATTERNS OF FISSURELLA SPECIES ON ISLA DE MARGARITA, VENEZUELA: USE OF RADULAE AND FOOD PASSAGE RATES

Structural differences and functional wear of the radula inthree species of the gastropod Fissurella from Isla de Margarita,Venezuela, were examined using light and electron microscopy.Wear patterns indicate between 6 and 9 transverse rows of teethare comonly used during feeding. Mechanical wear was most noticeableon the cusps of the outer lateral tooth; this wear varied fromrounding (F. nimbosa) to blunting (F. barbadensis) to squaring(F. nodosa) of the cusps. Morphological changes were additionallycharacterized by a significant decrease in the cusp length ofmarginal cusps in F. nodosa and breakage of the central toothand inner lateral teeth in F. barbadensis. Interspecific differencesin wear patterns suggest that the rhipidoglossate radula maybe used differently by congeneric Fissurella. Despite considerable variation, rasping rates while feedingon the same substrate were comparable among species; however,food passage rates through the digestive system differed amongspecies studied. Fissurella barbadensis requires 12 hours topass its food through the digestive tract, taking almost twiceas much time as F. nodosa and F. nimbosa. These data highlightdifferences in the feeding ecology of Fissurella species andcorrelate well with individual activity patterns and grazinghabits. ^*Present address: La Salle University, Department of Biology,20th Street at Olney Avenue, Philadelphia, PA 19141, USA. (Received 4 October 1988; accepted 16 February 1989)     

FOUR NEW SPECIES OF TAMBJA BURN, 1962 (NUDIBRANCHIA: POLYCERIDAE) FROM THE INDO-PACIFIC

The genus Tambja previously included 24 described species. Fournew species, T. tentaculata n. sp., T. gabrielae n. sp., T.zulu n. sp. and Tambja victoriae n. sp., from the Indo-Pacificare described. Tambja tentaculata n. sp., from Guam, is theonly known species in the genus with well developed, dorsolaterallygrooved, oral tentacles. Its inner lateral teeth have a bifidinner cusp with two long, sharp denticles. The oral tentaclesof T. tentaculata are more typical of Roboastra species, whilethe shape of the inner lateral teeth is more typical of Tambja.Nevertheless, the arrangement of the two cusps of the innerlateral teeth and the presence of a rachidian tooth withoutdenticles and with a central notch at the anterior edge, typicalof the species of the genus Tambja, suggest the placement withinthis genus. Tambja gabrielae n. sp., from Indonesia and PapuaNew Guinea, has dark green to dark brown ground colour withbright yellow patches scattered on the body. Tambja zulu n.sp. from Durban, South Africa, is characterized by a black groundcolour with slender yellow longitudinal lines. Tambja victoriaen. sp. is a new species from Papua New Guinea and Australiathat has frequently been misidentified as Roboastra arika, characterizedby its blue body colour and yellow lines. The four species aredistinguishable based on differences in body coloration, ofcharacters of the radula and of the reproductive system. Anoverview on distinguishing features of all known Indo-PacificTambja species is presented. (Received 21 June 2004; accepted 20 January 2005)