Variation in adult shell morphology and life-history traits in the land snail Oreohelix cooperi in relation to biotic and abiotic factors

In this study, we explore factors influencing variation in adultshell morphology and life-history characteristics (offspringsize and number) in the ovoviviparous land snail, Oreohelixcooperi, from the Black Hills, South Dakota and Wyoming, USA.To date, no study of shell morphology and life-history characteristicsin ovoviviparous land snails has included a combination of datasetsincluding genetic data, life-history traits, shell morphologyand multiple environmental factors. We report that differencesin shell size among populations are strongly related to meanannual temperature (and the highly correlated variable elevation)and population density (measured as shell density). In addition,shell size varies among populations, despite an apparent lackof population genetic differentiation. Common factors thoughtto influence adult shell size, like precipitation and calciumlevels, do not have a significant effect in this study. Adultsize strongly influences per-clutch reproductive output, withlarger snails having larger and more offspring. As mean annualtemperature and shell density affect adult shell size, theyalso indirectly affect per-clutch output. The results suggestthat a large portion of the life-history variation in O. cooperiis environmentally induced, as has been found in oviparous landsnails and brooding freshwater bivalves. (Received 20 July 2006; accepted 10 January 2007)     

The Production of Plantlets from Tissue Cultures of Brussels Sprout (Brassica oleracea L. var gemmifera D.C.)

Shoots were induced on callus derived from sprout sections andpetiole slices of an inbred parent line of Brussels sprout (Brassicaoleracea L. var gemmifera D.C.). The shoots, when excised andtransferred to fresh medium, enlarged and formed roots. Theseplantlets could be transferred to soil or their number increasedby a multiplication process involving the production of newshoots from the dormant lateral buds. Some of the plantletsderived from sprout callus were grown to maturity in the fieldand their morphology and chromosome number compared to seedgrown plants. There were no significant differences in sproutsize and stem diameter but there were significant differencesin plant shape. None of the plants in the field experiment showedpolyploidy. Plants derived from callus possessed an enhanced ability toform callus and redifferentiate when sections from these plantswere placed back on to nutrient medium.     

THE RELATIONSHIP BETWEEN SHELL INJURIES, SHELL THICKNESS AND HABITAT CHARACTERISTICS OF THE INTERTIDAL SNAIL LITTORINA RUDIS MATON

The hypothesis that risk of shell injury from crab predationand crushing by boulders and stones is important in the selectionof shell thickness in the intertidal gastropod Littorina rudisMaton was tested using data from 24 populations from Britain.Although there was an obvious relationship between the incidenceof shell injuries and both the distribution of crabs and substratummobility, there was no correlation between risk of shell injuryand shell thickness. This is probably accounted for by between-habitatvariations in energy levels which may vary independently ofcrab density and substratum mobility. ^*Present address: Department of Biology, University of Essex,Wivenhoe Park, Colchester C04 3SQ (Received 23 May 1977;     

CORRELATIONS BETWEEN SHAPE, WEIGHT AND THICKNESS OF SHELL IN FOUR POPULATIONS OF VENERUPIS RHOMBOIDES (PENNANT)

Three populations of Venerupis rhomboides in the Plymouth regionand one from the Firth of Clyde show remarkably wide, overlappingranges of variation in shape of shell. Shells of each populationare defined statistically in terms of relationships of shelllength, convexity and mean shell thickness, the latter measuredby a new technique utilizing sections of the shell cut radiallyfrom the umbo. Differences in shell shape are considered againstthe differences in the environments of the four populations,although environmental factors are incompletely known. The results of a transplantation experiment with mature bivalvesare also presented. The survivors showed correlated changesin shell shape and weight in the same direction and with thesame order of magnitude as those found in the distribution ofthe four populations. Detailed examination of the microscopicgrowth line patterns within the shells suggests that the increasedheight/length and convexity/length ratios and increased meanshell thickness were the results of reduced mantle exposureat the shell periphery in conditions of high concentration ofsuspended silt and clay. (Received 12 September 1982;     

Somatic Embryogenesis and Plant Regeneration from Inflorescence Culture of Java Citronella: (Cymbopogon winterianus)

Embryogenic callus was induced from immature inflorescence segmentsof Java citronella (Cymbopogon winterianus) and maintained for2 years on Murashige and Skoog's medium supplemented with 2,4-D(l mg l^–1). The callus cells retained the original chromosomenumber of 2n = 20. The somatic embryos germinated into plantletson MS basal medium or medium with IAA, NAA, BAP or KN individually(l mg l^–1). The regenerated plantlets developed a goodroot system on full strength solid MS inorganics medium withIAA (1 mg l^–1). The regenerated plants were similar tothe donor plant in morphology and had the same chromosome number,but showed some variation in the essential oil content. Java citronella, Cymbopogon winterianus, somatic embryogenesis, regeneration, inflorescence culture     

RELATIONSHIPS BETWEEN SHELL SHAPE, WATER RESERVES, SURVIVAL AND GROWTH OF HIGHSHORE LITTORINIDS UNDER EXPERIMENTAL CONDITIONS IN NEW SOUTH WALES, AUSTRALIA

The shell morphologies of the highshore littorinids, Littorinaunifasciata Gray and Nodilittorina pyramidalis (Quoy & Gaimard)have previously been shown to vary at a variety of spatial scales,including among replicate sites at the same height, from heightto height and from shore to shore. In this study, the relationshipsbetween morphology of the shell, the reserves of water heldwithin the shell, the size of the foot and survival on differentshores and rates of growth in different habitats were examinedfor L. unifasciata and, to a lesser extent, N. pyramidalis.Reserves of water were not consistently related to size or shapeof the shell, but did increase as relative weight of shell increased.This may be due to the relatively smaller body providing moreinternal volume for extra-corporeal water. Water reserves andthe amount of free water held in the shell were also not relatedto loss of water or survival during extended periods of emersion.Although the shape of the shell on sheltered and exposed shoreswas correlated with size of the foot, with the snails on anexposed shore having larger apertures and feet than those ona sheltered shore, transplant experiments did not show differentialmortality between morphs from the different shores. All translocatedand transplanted snails disappeared from the exposed shore ata greater rate than from the sheltered shore, but this was probablydue to the snails dispersing out of the experimental areas ratherthan due to mortality. Therefore, many of the large-scale modelsthat have previously been used to describe patterns of shellshape in intertidal gastropods do not appear to be importantin these highshore littorinids. Finally, field experiments ongrowth of juvenile L. unifasciata indicated that rate of growth,largely governed by opportunity to feed rather than type andquantity of food, is the most likely explanation for the small-and large-scale patterns of shell shape that have been previouslydescribed in this species. (Received 22 October 1996; accepted 19 February 1997)     

INTER AND INTRA-SPECIFIC VARIATION OF THE SHELL OF TWO SPECIES OF THE TROPICAL FRESHWATER SNAIL BELLAMYA (MOLLUSCA: PROSOBRANCHIA)

Shell variation in two species of the tropical freshwater snailBellamya from two different habitats, was studied. The variation of the shell features studied, which includedthe shell width, spire height, aperture height and aperturewidth, were found to be related to shell height. Furthermore,significant difference in the shell shape was found not onlybetween the two species but also between males and females ofeach species. (Received 18 August 1983;     

Ontogeny of the Behavior of Strombus maculatus (Gastropoda: Strombidae)

The ontogeny of strombid behavior was studied by observing thebehavior of Strombus maculatus veligers collected from the planktonand reared past metamorphosis to adults, and by observing juvenilestrombids collected in the field. Complete adult modal actionpatterns (MAP's) associated with locomotion, feeding, and rightingof overturned shells are performed by S. maculatus juvenilesimmediately after metamorphosis. There are changes in the frequencyof the use of certain MAP's which are associated with variationsin shell shape and size. The unique strombid escape response to molluscivorous gastropods(Conns spp.) is not released until juvenile S. maculatus arethree weeks past metamorphosis and two millimeters in shelllength. At that stage, the complete response is released uponthe first encounter with a predator. Experience with a predatordoes not seem to lower the age or size criteria. During ontogeny there is a trend toward an increasing complexityof behavior which is paralleled by an increasing complexityof neural structure and general morphology. There are majorsteps in the ontogeny of strombid behavior which probably coincidewith neural and morphological stages.     

Effect of Applied and Endogenous Auxin on Callus and Root Formation of In Vitro Shoots of the Apple: Rootstocks M26 and A2

The influence of exogenous IBA (indol-3yl-butyric acid) on rootand callus formation was studied in shoots of the apple rootstocksA2 and M26. The shoots grown in vitro were derived originallyfrom meristems of both juvenile and adult trees. Endogenousindol-3yl-acetic acid (IAA) concentrations in leaves and stemswere correlated with the responses to applied IAA. After 30 subcultures shoots from A2 and M26 rooted easily, butA2 did so more readily and even without IBA. Treatment withIBA improved percentage rooting and number of roots in bothrootstocks. Ex-adult and ex-juvenile shoots of A2 formed rootsto the same extent. However, ex-adult shoots of A2 showed ahigher IBA optimum for root number than ex-juvenile A2 and werealso less sensitive to supra-optimal IBA concentrations. Incontrast, in M26, there were no differences between ex-adultand ex-juvenile shoots. The results imply that rooting ability is associated more withdifferences between cultivars than with the origin of the explants.The best rooting occurred in ex-adult shoots of A2 which hadthe lowest endogenous IAA concentration, while callus formationwas correlated with high endogenous auxin concentration. Ex-adultA2 produced almost no callus even after exposure to high IBAconcentrations (25µM) whereas ex-adult M26 formed muchmore callus at 1/10 of the IBA concentration. Malus sylvestris (L.) Mill. var. domestica Borkh., Malus pumila Mill., apple rootstocks A2 and M26, in vitro culture, root and callus formation, HPLC analyses of IAA     

The Morphology, Growth and Reproduction of Unionidae (Bivalvia) in a Fenland Waterway

The shell morphology and population dynamics of the five BritishUnionidae are compared within a sympatric population. Pseudanodontacomplanata is distinguished from Anodonta anatina and A. cygneaby the hinge length–shell length relationship; this morphologicaldistinction may serve as a useful tool in the identificationof this threatened species. The shell length at a given annuluswas remarkably similar for all five species, although the asymptoticlength is reached most quickly in P. complanata and Unio pictorum.P. complanata is relatively short-lived and attains the lowest maximumlength, while A. cygnea lives more than twice as long and attainsalmost double the length of P. complanata. Unio spp. have ashort gravid season over the summer, while Anodonta spp. havea long gravid period, lasting from Autumn through to Spring.Unlike other members of the Anodontinae, P. complanata has ashort breeding season, overlapping with that of the Unio spp. (Received 4 March 1998; accepted 23 April 1998)     

SIZE, SHAPE AND SHELL MORPHOLOGY IN THE ANTARCTIC LIMPET NACELLA CONCINNA AT SIGNY ISLAND, SOUTH ORKNEY ISLANDS

Comparison of the size, shape and shell morphology in littoraland sub-littoral morphs of the Antarctic limpet Nacella concinnareveal differences in shell morphology which are enhanced bystructural anomalies within the shells of the two types. Infestationof sub-littoral shells by the conchocelis phase of an endolithicalga significantly affects shell density and total chlorophylllevels in the two shell morphs. The surface sculpture of sub-littoralshells is characterised by a series of grooves, the configurationof which closely resembles that of the radular teeth in N. concinna.Limpets utilise the available food supply within the shell matrixof other limpets by grazing the shell material. Epibiotic growthof calcareous algae prevent erosion and preserve underlyingshell layers. In severe cases, where protection is lacking,intraspecific shell grazing may remove parts of the shell exposingthe internal tissues. The Dominican Gull, Larus dominicanus, is a major shore predatorof both shell morphs. Gull middens contain both shell typesbut are dominated by the more accessible littoral shells. Comparisonof living populations and midden assemblages indicates thatsize and shape selection of prey occurs, with pear-shaped limpetsbetween 21 mm and 29 mm in length being taken preferentially. Apparent differences in shell form are induced by physical,biological and behavioural influences. Littoral animals arerobust in nature, resist avian pre-dation and are not extensivelygrazed whereas those of the sub-littoral are not subject tothe same degree of predatory attention but suffer a gradualdepletion of their shallower shell form through a combinationof algal infection and intraspecific shell grazing. (Received 21 February 1990; accepted 5 July 1990)     

Ribosomal RNA Gene Redundancy in Juvenile and Mature Ivy (Hedera helix)

Two-fold variation was found in rRNA gene redundancy in differentpopulations of ivy (Hedera helix). No consistent differencein rRNA gene number was observed between the juvenile and matureforms of the plant. Small significant differences between therRNA gene content of mature and juvenile forms and also betweenundeveloped buds and expanded leaves were found but these werein different directions in different populations.     

Detecting shape differences in species of the Littorina saxatilis complex by morphometric analysis

We investigated variation in shell size and shape of marinesnail species of the Littorina saxatilis complex (L. saxatilis,L. compressa and L. arcana) using geometric morphometric methods.These morphologically similar periwinkle species that are commonin the European intertidal have presented many problems fordiagnosis based on morphology alone. A discriminant analysisdemonstrated that geometric morphometrics is very efficientfor diagnosing individuals to species among sympatric populations.We successfully diagnosed an average of 96% of the specimens(with 85.7–100% correct diagnosis for specific comparisons).The diagnosis capability of this method is absolute at the populationlevel. This makes the technique potentially useful for the designof manipulative field or laboratory experiments. Moreover, ageometric-morphometric analysis was also accomplished in twosnail ecotypes (H and M) of L. saxatilis from rocky shores ofNE England which are apparently adapted to different degreesof wave exposure. We found that the H (exposed) ecotype hasa relatively rounded shell shape with a bigger aperture, whereasthe M (protected) ecotype has a smaller aperture. (Received 23 March 2006; accepted 25 January 2007)     

Within- and among-lake variation in shell morphology of the freshwater clam Elliptio complanata (Bivalvia: Unionidae) from south-central Ontario lakes

To compare the relative magnitude of variation in shell morphology within and among lakes, Elliptio complanata were collected from low and high exposure areas in each of four small lakes in south-central Ontario. Nested ANOVA's on shell length, height, width and weight revealed that shell morphology varied much more between sites of differing exposure within a lake than among lakes of differing alkalinity. Canonical variates analysis showed that clams from high exposure areas had larger and proportionately taller and heavier shells than those from low exposure areas. There was no relationship between alkalinity of lakes and shell morphology. These results suggest that the use of unionid shell morphology to predict long-term whole lake water chemistry (e.g. alkalinity) requires sampling designs which take into account within-lake variation in shell morphology.     

THE OPERCULA OF NERITOPSID GASTROPODS AND THEIR PHYLOGENETIC IMPORTANCE

The fossil record of neritopsid opercula and shells shows thatthe shell shape typical for Neritopsidae and Neritidae appearedin the Triassic. The ancestors of Neritimorpha were most probablyforms similar to Naticopsis. The operculum of Recent Neritopsisis composed of two calcitic parts secreted from inside and anaragonitic callus deposited from outside. Similar neritopsidopercula were already present in the Late Triassic. The firstopercula with asymmetrically situated muscle scars, possiblyancestral for neritids, also appeared at that time. (Received 27 May 2004; accepted 8 December 2004)     

SPATIAL PATTERNS OF SHELL SHAPE OF THREE SPECIES OF CO-EXISTING LITTORINID SNAILS IN NEW SOUTH WALES, AUSTRALIA

The shape and relative weight of the shell have been shown tovary intraspecifically and interspecifically in a number ofspecies of gastropods, including many different littorinids.These differences give rise to different shell forms in differenthabitats. In those species which have non-planktotrophic development,differences in shell form among shores have been usually explainedin terms of natural selection because exposure to waves supposedlyfavours light shells with large apertures, while predation bycrabs on sheltered shores favours elongated, thick shells withsmaller apertures. Differences in shell shape among speciesfound at different heights on the shore have been explainedin terms of resistance to desiccation and temperature. Suchvariables would tend to act on a relatively broad-scale, i.e.causing differences among heights on a shore or among shores.Rates of growth, which might vary at much smaller scales withina shore, have also been shown to affect the shapes of many shells. In this study, the shape and relative weight of shells of threespecies of co-existing littorinids (Littorina unifasciata, Bembiciumnanum and Nodilittorina pyramidalis) were measured. These speciesall haveplanktotrophic development and they are found on manyshores where there is no evidence that they are preyed uponby crabs. Before explanations of shell shape are proposed, itis necessary that patterns of variation, within different partsof ashore and among different shores are clearly documented.These patterns were measured at a number of different spatialscales within and among replicate shores with different amountsof wave exposure. Large and small specimens were included toallow intraspecific comparisons among snails of different sizesfound at different heights on the shore. The results showedsignificant differences among shores in shape and relative weightof shells, but these differences could not be explained by exposureto waves. In addition, snails of different sizes and differentspecies did not show the same patterns although they were collectedfrom the same sites. Importantly, the shell shape of Liuorinaunifasciata varied significantly among sites at approximatelythe same height within a shore. These differences could notbe clearly correlated with density, mean size nor exposure towaves. The only consistent pattern was a decrease in relativeaperture size in specimens living higher on the shore. Modelsthat have commonly been proposed to explain shape and relativeweight of shells in other species of gastropods are not adequateto explain the small- and large-scale variation of the measurementsdescribed here. It is proposed that any selective advantageof shell morphology and the effects of any variables on thedevelopment of shell morphology in these species can only beidentified after appropriately designed and replicated fieldexperiments. (Received 4 March 1994; accepted 13 September 1994)     

PREDATOR-PREY INTERACTIONS BETWEEN LEPSIELLA (BEDEVA) PAIVAE (GASTROPODA: MURICIDAE) AND KATELYSIA SCALARINA (BIVALVIA: VENERIDAE) IN PRINCESS ROYAL HARBOUR, WESTERN AUSTRALIA

On a sandy beach at Shoal Bay in Princess Royal Harbour, Albany,southwestern Western Australia, lives a small muricid gastropodthat feeds virtually monotonically on the overwhelmingly dominantresident bivalve Katelysia scalarina. Lepsiella paivae livesburied in the sand and attacks its prey within it. Because ofits small size (<13 mm shell height), bivalve prey isalso small and this study demonstrates a preference for K. scalarinaof 5 mm shell length, i.e. juveniles. Laboratory experimentsalso suggested a possible preference for attack of the rightvalve. Lepsiella paivae can and does, however, attack largerprey (up to 15 mm shell length), but cannot consume themcompletely. A second visit to Princess Royal Harbour in theAustral winter, when there was no juvenile K. scalarina present,showed L. paivae to be attacking at the sand surface, also bydrilling, the small (<4 mm) gastropod Hydrococcus brazieri(Hydrococcidae). SEM studies of experimentally determined drillholes of L. paivae show them to be of variable form, some straightsided, others bevelled (like a naticid) and <500 µmin diameter. On this sheltered Southern Ocean beach, therefore,L. paivae has specialized to attack juvenile bivalves by burrowingafter them. It can, however, attack other species opportunisticallyon the sand surface when seasonally favoured juvenile bivalveprey are not present. (Received 8 January 2005; accepted 16 March 2005)     

Variation in shell morphology of Calliostoma zizyphinum (Gastropoda: Trochidae)

Differences in shell morphology in the intertidal prosobranchmollusc Calliostoma zizyphinum were studied from a number ofsites within four geographical regions of the British Isleswith varying exposures to wave action and crab predation. Meanvalues of damage scarring were highest in shells sampled fromStrangford Lough, Northern Ireland, and lowest in individualsfrom the Atlantic Coast. Shells collected from the Isle of Manwere smaller than those from either the Atlantic coast of Irelandor Strangford Lough. Shells from Strangford Lough had tallershells (higher aspect ratio) than shells from the County Downand Atlantic Coast and shells from the County Down coast weremore squat (lower aspect ratio) than those from all other areas.Shells from Strangford Lough were significantly thinner thanshells from all other geographical areas. The relationshipsbetween shell damage scarring and shell size and tallness werenot consistent among areas. Shells from Strangford Lough arenotable in showing a steep, positive relationship between shellsize and scarring and a steep, negative relationship betweenscarring and tallness. The unusual shell phenotypes observedin shells from Strangford Lough may be explained by rapid shellgrowth, which would not only allow Calliostoma to attain a sizerefuge from crab predators but also to recover successfullyfrom repeated crab attacks on the shell lip. Such a strategywould result in larger, thinner shells with a high number ofdamage scars. (Received 6 June 2006; accepted 20 December 2006)     

INTRA-POPULATION VARIATION OF THE SHELL OF LITTORINA RUDIS (MATON) (MOLLUSCA: PROSOBRANCHIA)

Shell variation within a single population of Litlorina rudis(Maton), collected near the Biological Station at Trondheim,Norway, was studied. The variation of the shell features studied, which includedthe shape of the basal part of the outer lip, and the relativeheight of the spire, width of the shell and of the aperture,were found to be at least partially related to shell height.Furthermore, the shell shape in the population investigated,besides varying with age, also varies due to shell damage. Nosignificant differences were found between the shapes of theshells of males and females. The shell characteristics of different populations of L. rudisalso vary greatly, one of the causes being differences in theenvironmental conditions. When studying this variation betweendifferent populations, however, it must be emphasized that onlyundamaged shells of individuals of about the same age shouldbe compared, in order to obviate the effects of the variabilitydue to age differences and shell damage within each individualpopulation. ^*Contribution from Trondhjem Biological Station no. 199 (Received 18 February 1980;     

GEOGRAPHIC VARIATION AND HETEROCHRONY IN TWO SPECIES OF COWRIES (GENUS CYPRAEA)

Geographic variation in the marine, Indo-Pacific cowry, Cypraea caputserpentis, involves clinal variations that parallel the ontogenetic development of adult shell characteristics. Cypraea caputdraconis, a closely related species endemic to Easter Island and Sala y Gómez, is morphologically similar to juvenile C. caputserpentis. Using multivariate measures of size and shape, I examine these patterns as a possible outcome of heterochrony, or changes in the timing of developmental events in ontogeny. Whorl-expansion rates of juvenile shells are significantly higher in C. caputdraconis when compared to C. caputserpentis and are negatively correlated with surface seawater temperatures among populations of C caputserpentis. High expansion rates, often associated with slow growth, result in a delay in the onset of lateral callus development and subsequent paedomorphosis. Ontogenetic trajectories calculated from growth series of adult and preadult shells indicate that paedomorphosis results from the combined effects of neoteny and post-displacement. Paedomorphosis among cowries may result from the advantages of larger body size relative to shell size under reduced predation intensities and associated increases in fecundity.