Molluscan Metamorphosis: A Study in Tissue Transformation

Metamorphosis of the pelagic larvae of benthic marine invertebratesis often a cataclysmic event in which a rapid loss of organsspecialized for larval life occurs simultaneously with the renewalor increased rate of development of potential adult organs.In the nudibranch gastropod Phestilla sibogae this change involvesloss of the velum, shell, operculum, larval kidney, some retractormuscles, and some of the pedal mucous glands. Exit from thelarval shell at metamorphosis is rapid and is correlated withthe spread of epidermis from the larval foot over the visceralmass as the visceral mass emerges from the shell aperature.This spreading of epipodial epidermis to cover the entire bodyhas not been previously reported for other nudibranchs. Neithercell proliferation nor active cell motility are responsiblefor this epidermal migration. Rather it appears that the actionof larval muscles pulls the visceral organs out of the shelland simultaneously causes the epipodial epidermis to cover thevisceral mass. This epidermis becomes the definitive adult epidermis.     

Systematics and Ecology of Shell-Boring Polychaetes from New England

In a survey of shell-boring polychaeles of New England, representativesof five families were found: Spionidae, Cirratulidae, Capitellidae,Terebellidae, and Sabellidae. The five spionid species were studied extensively. These includePolydora commensalis Andrews, found only in shells occupiedby hermit crabs, P. concharum Verrill, P. socialis (Schmarda), P. websleri Hartman, and Boccardia hamata (Webster) , foundin various types of shell. Breeding periods were defined andthe larval development described for each species. Polydoraconcharum deposits egg capsules inwinter months, while the oilierpolydorids spawn in spring or summer. Morphology of planktoniclarvae is distinct in each species, although P. websteri larvaemay be confused with non-boring species such as P. ligni Webster.Polydora socialis adults are found in both shells and sedimentand have a unique grinding apparatus, a gizzard, between theesophagus and intestine. The cirratulid, Dodecaceria sp., follows an asexual mode ofreproduction. The syslematics of Dodecaceria is complex owingto multiple modes of reproduction. No sexually mature individualswere observed during the course of this study. Asexual budswerefound in the Fall. A sabellid, Pseudopolamilla reniformis (Miiller), is commonlyfound in theshells of Placopeclen magellanicits (Gmelin) inMaine waters. Its reproductionis unknown.     

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)     

Seasonal and tidal abundance patterns of decapod crustacean larvae in a shallow inlet (SW Spain)

Planktonic larvae of decapod crustaceans were collected monthlyfrom July 1991 to June 1992 by pumping during nocturnal floodand ebb tides to establish seasonal larval abundance patternsin an inlet of the Bay of Cdiz. Additional 24 h series of sampleswere collected seasonally (July 1991, October 1991, January1992 and May/June 1992) during spring and neap tides to analyselarval abundance in relation to the main environmental cycles(diel, tidal and lunar phases) and vertical position in thewater column. First zoeae were the most abundant stage for mostspecies, representing 97.6% of all individuals collected. ZoeaI abundance was higher in spring and swmner and, on most samplingoccasions, there was a net output from the inlet to the bay.Five species (Liocarcinus arcuatus and Liocarcinus vernalis,Uca tangeri, Diogenes pugilator and Panopeus africanus) represented60% of total individuals caught. The seasonal occurrence offirst zoeae of the most abundant species indicated two differentreproductive patterns: species with a short reproductive periodand species spawning year round. Zoea I of several species (Panopeusafricanus, Uca tangeri, Pachygrap sus marmoratus, Processa spp.)were significantly more abundant during ebb tides and theirlater larval stages were scarcely collected, suggesting thatthese larvae are released in the inlet and exported to the bay.Conversely, a net input of first zoeae was observed for otherspecies (D.pugilator and Pinnotheres pinnotheres), but theirlater larval stages were also scarcely collected. Such importationcould be a larval rhythm artifact due to release of larvae inthe bay that drifted into the inlet by tidal currents. The crabIlia nucleus, whose later larval stages were collected frequently,was the only species that seemed to complete its life cyclewithin the bay. These results suggest that the studied inletwas primarily used by decapods as an adult habitat and spawningground, while larval development occurred in open sea. Sincevertical migration was not observed for exported larvae, thetidal synchronization of female release seemed to be the mostprobable mechanism of larval exportation. There were no significantdifferences between larval release during spring and neap tides.     

棉铃虫对三氟氯氰菊酯的抗性狭义遗传力及其与体重的相关性

利用数量遗传学方法和半同胞交配设计,测定了棉铃虫Helicoverpa armigera对三氟氯氰菊酯抗性和体重的狭义遗传力,并分析了棉铃虫抗药性与其体重之间的相关性。结果表明,棉铃虫对三氟氯氰菊酯的抗性狭义遗传力为0.2476±0.0248,体重遗传力为0.3613 ± 0.1299;抗性与其母体效应无关;抗性与体重之间存在显著的遗传负相关。     

Embryonic and larval development of Ensis arcuatus (Jeffreys, 1865) (Bivalvia: Pharidae)

The razor clam Ensis arcuatus (Jeffreys, 1865) is distributedfrom Norway to Spain and along the British coast, where it livesburied in sand in low intertidal and subtidal areas. This workis the first study to research the embryology and larval developmentof this species of razor clam, using light and scanning electronmicroscopy. A new method, consisting of changing water levelsusing tide simulations with brief dry periods, was developedto induce spawning in this species. The blastula was the firstmotile stage and in the gastrula stage the vitelline coat waslost. The shell field appeared in the late gastrula. The trochophoredeveloped by about 19 h post-fertilization (hpf) (19°C).At 30 hpf the D-shaped larva showed a developed digestive systemconsisting of a mouth, a foregut, a digestive gland followedby an intestine and an anus. Larvae spontaneously settled after20 days at a length of 378 µm. (Received 5 December 2006; accepted 19 November 2007)     

Larval shells of Late Palaeozoic naticopsid gastropods (Neritopsoidea: Neritimorpha) with a discussion of the early neritimorph evolution

Extant neritimorphs with planktotrophic larval development have a convolute smooth larval shell which is internally resorbed. The oldest known larval shells of this type are of Triassic age. Well-preserved Late Palaeozoic neritimorph specimens have larval shells of two or more rapidly increasing well separated whorls. These larval shells resemble planktotrophic caenogastropod larval shells. This type of larval shell is possibly plesiomorphic in neritimorphs and caenogastropods. Permian/Pennsylvanian neritimorphs (Naticopsis, Trachyspird) have smooth larval shells (Naticopsidae) or larval shells with strong axial ribs (Trachyspiridae new family). The convolute low-spired round shell shape of modern neritimorphs is causally linked with the resorption of the inner teleoconch and protoconch whorls. Modern neritimorph shells with a uniform, undifferentiated inner lumen have probably evolved from naticopsid ancestors which lack resorption. It is possible that an elevated spire, deep sutures and protruding spiral larval shells would have made such internally undifferentiated shells more vulnerable for mechanical destruction and prédation. Suggestions that coiling evolved independently in neritimorphs and other Gastropoda are unlikely and contrast with the fossil record. The modern neritid larval shell has probably evolved from relatively low-spired smooth naticopsid larval shells like those reported here.     

Effects of temporary food limitation on survival and development of brachyuran crab larvae

As a consequence of the combined effects of prey patchinessand diel or tidal vertical migrations in the water column, decapodcrustacean larvae may experience temporal or spatial variabilityin the availability of planktonic food. In a laboratory study,we evaluated effects of temporarily limited access to prey onthe larvae of three species of brachyuran crabs, Chasmagnathusgranulata, Cancer pagurus and Carcinus maenas. Stage-I zoeaewere fed ad libitum for 4 or 6 h per day (20 or 25% treatments;6 h tested in C. pagurus only), and rates of larval survivaland development were compared with those observed in continuouslyfed control groups (24 h, 100%). In C. granulata, we also testedif intraspecific variability in initial biomass of freshly hatchedlarvae originating from different broods has an influence onearly larval tolerance of food limitation. Moreover, we exposedembryos and larvae of this estuarine species to moderately decreasedsalinities to identify possible interactions of osmotic andnutritional stress. Finally, we evaluated in this species theeffect of food limitation on survival from hatching throughall larval instars to metamorphosis. In all three species, limitedaccess to prey had only weak or insignificant negative effectson survival through the Zoea-I stage. The strength of the effectsof temporary food limitation varied in C. granulata significantlyamong broods. However, no significant relationships were foundbetween initial larval biomass (C content) and either survivalor development duration. Strongly decreased survival to metamorphosiswas found when food limitation continued throughout larval development.Thus, early brachyuran crab larvae are well adapted to transitorylack of planktonic food. The capability of the Zoea-I stageof C. granulata to withstand nutritional stress also under conditionsof concomitant salinity stress allows them to exploit variousbrackish environments within estuarine gradients. However, continuedexposure to limited access to planktonic prey may exceed thenutritional flexibility of C. granulata larvae.     

IN VITRO RECALCIFICATION OF ORGANIC MATRIX OF SCALLOP SHELL AND SERPULID TUBES

Organic matrix was isolated from the shell of the bivalve Argopectenirradians by decalcification. The capacity of the matrix toinitiate formation of crystals similar in form and orientationto the crystals of normal shell was investigated. Decalcifiedshell matrix placed in an inorganic recalcification solutioninitiated the formation of elongate crystals in parallel arrangementcorresponding to the parallel orientation observed in the matrixfibers and similar to the orientation of the long crystals innormal shell. The detailed form of the crystals deposited invitro was different from that of the normal shell crystals.Electron diffraction analysis of remineralized matrix demonstratedthat the material was calcite, the mineral of normal shell. In contrast, the calcareous tube of the serpulid Hydroides dianthushas crystals lacking uniform arrangement and a matrix whichdoes not have a well-oriented structure. The decalcified tubematrix was recalcified and the mineral posited showed some evidenceof normal orientation. The results demonstrate that matrices of Argopecten shell andHydroides tube can induce crystal formation in vitro. Sincethe soluble matrix would be expected to be removed during decalcification,the observed in vitro effects apparently involve the insolublematrix. (Received 19 June 1984;     

CONVERGENT SHELL MORPHOLOGY IN INTERTIDAL GASTROPODS

The functional morphology of shell infrastructure in 2 speciesof intertidal trochid was compared with that in 2 species ofnerite. The shell of Monodonta constrictais typical of the majorityof trochids. The shell is composed of 4 layers: a distal layer(calcite), anouter prismatic layer (aragonite), a nacreous layer(aragonite), and an oblique prismatic layer (aragonite). Monodontalabio lacks a distal layer and an oblique prismatic layer. Theoblique prismatic layer is replaced by an inner prismatic layerwhich forms an apertural ridge as a result of deposition andresorption. The shells of Nerita versicolor and N. tessellataconsistof 3 layers: an outer prismatic layer (calcite), a crossedlamellar layer (aragonite), and a complex crossed lamellar layer(aragonite). The complex crossed lamellar layer is covered withinclined platelets which superficially resemble the surfaceof the ique prismatic layer of trochids. In addition, the complexcrossed lamellar layer forms an apertural ridge which is similarin appearance to that of Monodonta labio. The outer surfaceof the mantle of Nerita versicolor and N. tessellata is throwninto a series of large folds which lie in contact with the inclinedplatelets of the omplex crossed lamellar layer. The interactionof the mantle folds with the inclined platelets is thought toserve as a rachet mechanism to aid in extension of themantle;a similar function has previously been proposed for trochids.The apertural ridges of Monodonta labio and Nerita are thoughtto prevent excessive desiccation when these gastropodsare exposedat low tide. ¹Contribution No. 56 of the Tallahassee, Sopchoppy & GulfCoast Marine Biological Association (Received 6 July 1979;     

INTRASPECIFIC AND INTERSPECIFIC VARIATION IN SHELL MORPHOLOGY OF BIOMPHALARIA PFEIFFERI (KRAUSS, 1848) AND HELISOMA DURYI (WETHERBY, 1879) FROM MOSHI, TANZANIA, ANALYSED BY MORPHOMETRIC STUDIES

Helisoma duryi, a planorbid snail species, which is a potentialbiological control agent of the intermediate hosts of schistosomiasis,was introduced in an irrigation scheme (T.P.C.) in northernTanzania in 1972. The species has been present at T.P.C. since,and during a snail survey in January 1981, a great variationin the shell morphology of H. duryi, ranging from typical H.duryi forms to forms resembling Biomphalaria pfeifferi, theintermediate host of Schistosoma mansoni in the area, was observed.The Biomphalaria- Wkc forms of H. duryi could constitute anentire population but also occasionally occurred in populationsof typical H. duryi. This study was undertaken to determine whether simple shellmeasurements, or ratios between some of these could be usefulin discriminating between H. duryi and B. pfeifferi. Followingparameters were considered: shell height, shell diameter, diameterof the umbilicus, ratio between shell height and diameter ofthe umbilicus, and the number of whorls. (Received 29 July 1983;     

ALTITUDINAL VARIATION AND ADAPTIVE ZONES IN ARIANTA ARBUSTORUM: A NEW LOOK AT A WIDESPREAD SPECIES

Arianta arbustorum is a European land snail, occurring in bothlowland and mountainous regions. Its shell is usually globular,which makes the species aberrant among the Campylaeinae in shellshape, habitat, and wide, not exclusively montane range. Withincreasing altitudes, the shells usually decrease in size andbecome more conical. However, in some mountain stocks thereare populations with depressed shells, provided with an openumbilicus. This form (subspecies) is only known from formerPleistocene refugia. It is hypothesized that in the Pliocene, the primary montaneA. arbustorum, with a depressed shell, entered an additional,adaptive zone, while conquering damp lowland habitats. Thisresulted in an enormous range extension. In the new habitata different (globular) shell shape evolved and the species becamepolytypic. The subsequent glaciations wiped out most of theoriginal montane A. arbustorum with depressed shells. Wherethis had occurred, post-Pleistocene re-colonizing conspecificsnails with globular shells, found vacant territory and coulddevelop the dines mentioned before. The invading populationsdid not regain the original, montane, depressed shell shape.This shape was only retained in a few refugia, where the populationssurvived amidst the glaciers. (Received 9 February 1990; accepted 31 May 1990)     

Naupliar development of Acanthodiaptomus denticornis (wierzejski, 1887) and Arctodiaptomus alpinus (Imhof, 1885) (Copepoda: Calanoida) and a comparison with other Diaptomidae

The six naupliar instars of the two alpine species Aconthodiaptornusdenticornis and Arctodiaptomus alpinus are described and illustrated.Their external morphology is compared with that of all presentlyknown diaptomid nauplii in an attempt to provide usehl taxonomiccharacteristics to identify larval diaptomids. The larval stagesof the two species are remarkably similar in size and overallappearance, and show an identical pattern of limb setation throughoutthe whole development. Diagnostic characters are mainly relatedto the differentiation of antennules and caudal armature.     

Natural variability in size and condition at settlement of 3 species of marine invertebrates

Experimental studies have demonstrated that for many marineinvertebrate species, variability in larval condition or qualityat settlement may have important effects on post-settlement,early juvenile performance. Relatively few studies, however,explicitly examine natural variability in larval condition atsettlement. This study examines natural variability in larvalattributes (size and lipid index) at settlement for terminal-stagelarvae of intertidal mussels (Mytilus sp.) and barnacles (Pollicipespolymerus and Chthamalus dalli) from southern California. Despitesignificant differences among cohorts in larval attributes,for all 3 species a greater percentage of the variance in larvallength (80–100%) and lipids (58–83%) occurred amongindividuals within a cohort, rather than among cohorts. Forall 3 species, coefficients of variation within a cohort forlength were much smaller (3–8%) than those for lipid index(30–93%), suggesting that lipid storage is a much moreplastic attribute than size for larvae. For mussels, settlementintensity and larval attributes were decoupled, such that averagelarval condition of a cohort was not related to the number oflarvae that settled. At the cohort level, Mytilus and Pollicipessettling together across 3 dates showed similar trends of decreasinglipid index over time, suggesting that environmental conditionsmay influence co-occurring planktonic larvae similarly acrossspecies. This work highlights the need for further experimentsin the field on the effects of larval history on recruitmentsuccess in natural populations, and further studies to determinewhat factors influence larval attributes for planktonic larvaein the field.     

Control and Consequences of Alternative Developmental Modes in a Poecilogonous Polychaete

SYNOPSIS. The poecilogonous polychaete Streblospio benedicti(Webster) exhibits both planktotrophic and lecithotrophic modesof larval development. The alternative trophic modes are associatedwith differences in age and size at maturation, offspring number,size and energetic investment, larval planktonic period, morphologyand survivorship. This paper reviews a decade of research intothe control and consequences of the traits associated with planktotrophyand lecithotrophy in S. benedicti. The dominant control on reproductiveand developmental characters is genetic. Significant additivegenetic variance has been detected for egg diameter, fecundity,larval planktonic period and aspects of larval morphology. However,environmental factors such as temperature, food quality andphotoperiod, and intrinsic factors such as maternal age, exertconsiderable influence on non-trophic developmental traits (e.g.,offspring number, size and energy content). Demographic consequencesof development mode are reviewed for field and laboratory demesof S. benedicti dominated by individuals exhibiting either planktotrophyor lecithotrophy. Similar population size structure, fluctuationsin abundance, P: B ratios, and estimated population growth ratesare achieved through trade-offs between survivorship and fecundity. Development mode may best be viewed as a complex set of traitsthat are intimately linked developmentally and evolutionarilyto other aspects of an organism's life history. Greater insightinto the control and consequences of development mode shouldresult from further investigation of these linkages     

Complete larval and early juvenile development of the mangrove crab Perisesarma fasciatum (Crustacea: Brachyura: Sesarmidae) from Singapore, with a larval comparison of Parasesarma and Perisesarma

The complete larval development of the sesarmid crab Perisesarmafasciatum (Lanchester, 1900) from Singapore was obtained fromlaboratory culture. All four zoeal stages, the megalopa andthe first crab stage are described and illustrated. The morphologicalcharacteristics of the larvae of P. fasciatum are compared withthose of other known larvae of the genera Perisesarma and Parasesarma.The larval morphology of P. fasciatum clearly presents the typicalcombination of features that characterize sesarmid larvae. Overall,larval stages are very similar in Perisesarma and Parasesarmaand it is impossible to distinguish these two genera by larvalmorphology.     

THE ROLE OF LYMNAEA AURICULARIA (LINNAEUS) AND LYMNAEA PEREGRA (MULLER) (GASTROPODA: PULMONATA) IN THE TRANSMISSION OF LARVAL DIGENEANS IN THE LOWER THAMES VALLEY

Eleven species of larval digeneans are reported for the firsttime in the UK from the freshwater snail Lymnaea (Radix) auricularia.The latter replaced its congener L. peregra in a gravel pitnear Wraysbury, Berkshire in the Lower Thames Valley and hostedan assemblage of larval digeneans similar to that previouslytransmitted by L peregra. The larval digeneans of L. peregrain small bodies of water in the area were also similar to thosefound in L. auriculania in the gravel pit. This replacementin the role of the host is explained on the basis of morphological,physiological and ecological similarities of the two lymnaeidspecies. (Received 19 February 1992; accepted 27 April 1992)     

Atebbania Bernasconii, a New Genus and Species from Subterranean Waters of the Tiznit Plain, Southern Morocco (Gastropoda: Hydrobiidae)

The new genus Atebbania is described for a new hydrobiid species,A. bernasconii, which lives in groundwaters of southern Morocco.Of the Hydrobiidae genera with known shell and anatomy Moitessieriais the closest to Atebbania. The two share the following characters:shell elongate; teleoconch surface with evident microsculpture;seminal receptacle absent. Atebbania is distinguished from Moitessieriaby: shell ovate; penis with a) apical stylet, b) one lobe bentdownward on left (inner) side and c) penial duct running nearthe right (outer) side; posterior end of foot indented. Atebbaniabernasconii n. sp., lives in a limited area, the Tiznit plainin southern Morocco. Other stygobiont hydrobiids from otherareas of Morocco are currently being studied. None of them appearsto be closely related to A. bernasconii or to the species ofMoitessieria (Received 10 February 1998; accepted 30 April 1998)