Encapsulated development of the marine prosobranch gastropod Nucella lapillus

Nucella (=Thais) lapillus (L.) develops to the juvenile stage within an egg capsule. This study was undertaken to investigate nutritive and bacteriostatic properties of the intracapsular fluid for this species. Capsules were collected intertidally at Nahant, Massachusetts, USA, in the summers of 1982 and 1983. The average capsule contained only 1.1 l of intracapsular fluid per embryo. A statistically significant correlation was observed between capsule height and number of developing embryos contained therein, although the number of embryos per capsule was more closely related to capsule volume. Total dry weight of encapsulated, shelled embryos increased exponentially as a function of shell length. However, weight increases may be entirely due to shell calcification; the average dry tissue weight of shelled embryos was significantly below that for pre-shelled individuals, suggesting a net loss of biomass during development. Aliquots of intracapsular fluid failed to inhibit the growth of the 13 bacterial strains tested. Despite the above results, pre-shelled embryos generally survived only several days following their premature removal from egg capsules at 12°C. Shelled veligers were reared outside of the capsules with no apparent ill effects. An explanation for these survivorship results is elusive.     

An alternative mechanism to reduce intracapsular hypoxia in ovicapsules of Fusitriton oregonensis (Gastropoda)

Oxygen supply inside egg masses has been reported as a constraint of embryo development. Many species that enclose their eggs in jelly masses or ovicapsules have strategies to avoid or reduce intracapsular hypoxia. In some amphibian species, a decrease in the wall thickness of the egg capsule over time produces an increase in oxygen conductance of the wall, reducing the problem of intracapsular oxygen limitation. Previous studies of gastropods have reported a decrease in the thickness of the capsules during development. However, there are no studies relating capsule thinning to the oxygen limitation problem in this group. This study links the thinning of egg capsules with oxygen diffusion as a possible mechanism to reduce or avoid hypoxia inside the capsules of the gastropod Fusitriton oregonensis. Capsule thickness, capsule area, oxygen partial pressure inside and outside the capsule, and oxygen consumption of the embryos at early and late developmental stages were measured. The conductance and the diffusion coefficients of the capsule were estimated using these measurements. Results showed that (1) capsule thickness decreased throughout development by about 50%, (2) oxygen consumption of embryos increased from early to late stages, (3) oxygen partial pressure inside the capsule did not change during development, (4) conductance coefficient increased with time, and (5) estimation of diffusion coefficient was lower than amphibian egg jelly, shark capsules, egg fishes, and eggs of giant cuttlefish. The reduction in the thickness of the capsule wall and the associated increase in its conductance during embryonic development may reduce oxygen constraints, especially at late developmental stages.     

Osmotic and ionic concentration of the egg capsule fluid of Crepidula fornicata in relation to embryonic development

Osmotic pressure and major ions (Cl⁻, Na⁺, Mg²⁺, Ca²⁺) of the egg capsule fluid in the slipper limpet Crepidula fornicata were investigated in relation to embryonic development. Calcium permeability of the capsule wall was studied at oviposition, by dipping freshly laid egg capsules in ⁴⁵Ca as a tracer. This study also determined total calcium content of the embryos at different developmental stages. Osmolarity and major ion concentrations in egg capsule fluid were higher than seawater at uncleaved and trochophore stages, and then dropped to the same level as sea water at veliger stage. Concentrations of Cl⁻ and Na⁺ were relatively high at oviposition, peaked at trochophore stage, and finally dropped close to concentrations of seawater at hatching. In contrast, concentrations of Mg²⁺ and Ca²⁺ decreased steadily during capsular development. Radiotracer permeability experiments in freshly laid egg capsules confirmed that the capsule wall is impermeable to this ion at that stage. However, because of the dissolution of the inner layer of the wall during the final part of capsular development, the wall becomes permeable to calcium and probably to the rest of the major ions studied.     

A correlated morphological and biochemical study of capsular fluid of Nucella lapillus (Gastropoda: Prosobranchia: Muricidae)

The extra-embryonic albuminous capsule fluid of the marine prosobranch Nucella lapillus L., collected from March to May 1988 in Brittany, France, has been investigated by sodium dodecylsulfate-polyacrylamide-gelelectrophoresis (SDS-PAGE) and biochemical assays. The fluid at the gastrula- and early veliger stage of the snail embryo was analysed for proteins on SDS-gels, i.e., for acid phosphatase, alkaline phosphatase, -glucuronidase and arylsulfatase activities. The protein and inorganic phosphorous contents were also determined. Electrophoresis revealed the presence of different proteins and a variation in the band pattern during early developmental stages. The total protein content of the capsule fluid decreases significantly from gastrula to early veliger stage. In the differentiated developmental stage the acid phosphatase activity was higher than the poorly differentiated gastrula stage. The detected activity of arylsulfatase indicates that acid mucopolysaccharides are present. In general, observed changes in the organic components of the intracapsular fluid of N. lapillus substantiate the nutritive value for developing embryos.     

Combined effects of dissolved oxygen concentration and water temperature on embryonic development and larval shell secretion in the marine snail <Emphasis Type="Italic">Chorus</Emphasis><Emphasis Type="Italic">giganteus</Emphasis> (Gastropoda: Muricidae)

The present study was undertaken to determine the effects of both extracapsular oxygen concentration and temperature on embryonic development in Chorus giganteus. In normoxia increasing water temperature from 12°C to 18°C reduced by 15 days the median time required for the capsules to hatch. Hypoxia (oxygen content at 50% of air saturation) generated a low development rate and totally prevented both shell secretion and larval hatching from the egg capsule. Experimental transfer at weekly intervals, from normoxia to hypoxia and vice versa, induced a decrease and increase in the embryonic ash content, respectively, but did not affect the number of hatched larvae. Such an effect was more pronounced at 12°C than at 15°C or 18°C. The embryonic inability to produce a shell under hypoxia is likely to be a result of the low intracapsular oxygen concentration (IPO₂) generated as the combined effect of a low extracapsular oxygen concentration (environmental) added to the intracapsular embryonic oxygen demands, which lowers the IPO₂ still further. Under such conditions, a decrease in intracapsular pH is likely to take place, and, if so, embryos might divert carbonates away from shell calcification to balance such changes in pH.     

Predation on egg capsules of <Emphasis Type="Italic">Zidona</Emphasis><Emphasis Type="Italic">dufresnei</Emphasis> (Volutidae): ecological implications

Among the diverse patterns of energy allocation to the offspring of gastropods, the presence of egg capsules to protect embryos is common. Females of the edible snail Zidona dufresnei attach egg capsules to hard substrates in shallow Argentine Patagonian waters (40°45′S, 64°56′W) during spring-summer. Embryonic development takes about 30 days at 22°C. In this study, three likely capsule predator species and the marks left by each on egg capsule walls were identified in laboratory experiments in February 2010. Abundances of predators and egg capsules with evidence of predation were assessed in the field in the summers of 2010 and 2011. Under laboratory conditions (N = 10 replicates per treatment and control), the predation rate by the chiton Chaetopleura isabellei was the highest (up to 90%), followed by the gastropod Tegula patagonica and the crab Neohelice granulata (~20% each). Nearly 60% of 41 capsules found in the field showed signs of predation. According to the marks identified in the laboratory, C. isabellei was responsible for 79% of this predation, and T. patagonica for the rest. Predation appears to be important during the encapsulated early life and could be an agent for selecting for resistant capsule walls and a relatively shorter development time.     

Embryo development and maternal–embryo nutritional relationships of piked spurdog (<Emphasis Type="Italic">Squalus</Emphasis><Emphasis Type="Italic"> megalops</Emphasis>)

The maternal–embryo relationship was determined for the piked spurdog (Squalus megalops). In addition, the increase in offspring size with maternal size was studied and the embryonic development was described. Wet weight of in utero eggs and offspring size was correlated with maternal size; larger females produced larger embryos which would have higher survival rate and reproductive value. All embryos present in a female were at a similar stage of development. The external yolk sac is reabsorbed late in gestation, suggesting that embryos are mostly nourished by yolk sac reserves. Embryo size-at-birth varied considerably (180–244 mm total length) as a result of the significant variability in ova size at ovulation. The amounts of water, organic and inorganic matter of embryos at different stages of development were measured to determine possible maternal contributions during embryonic development. Total wet weight from smallest and largest in utero eggs to smallest and largest term embryos changed by +46 and +58%, respectively. This pattern was due to a change in water content by +137 and +154%, and inorganic matter by +100 and +156%. Organic matter of smallest and largest in utero eggs changed by −23 and −17%, respectively. The uterus of pregnant females became specialised for water and mineral transport, not nutrient provision. These results indicate that S. megalops is a strict yolk-sac viviparous species with no maternal contribution of organic matter during development.     

Reproduction and sex reversal of the solitary vermetid gastropod<Emphasis Type="Italic"> Serpulorbis arenarius</Emphasis>

The reproductive biology of the Mediterranean vermetid gastropod Serpulorbis arenarius, a subtidal sessile solitary species that lives in shady rocky communities, was studied at two localities south-east of the Iberian Peninsula. Its population density is usually low, with specimens sometimes sparsely distributed. Its shell often appears to be covered by epibiotic organisms. The sex ratio is biased toward males (63%) in the populations of SE Spain, and the results point towards hermaphroditism with alternating sexuality. The results obtained suggest that all specimens are first males, change to females under some as yet unknown condition and, after the ovogenesis cycle has been completed, revert to the original sexual state (male). The possible adaptive forces that favour hermaphroditism in this species are discussed, and we hypothesise that the seizure of a spermatophore by males may stimulate sex change to female. Brooding females (51% of females studied) were found from March to December. In males, histological study of testes revealed that spermatogenesis seems to occur throughout the year. Females brooded between 5 and 28 (=15) egg capsules, which were attached by stalks to the inner surface of the shell. It is noteworthy that the gonads of two of the specimens brooding egg capsules were developing testes. The newly laid egg capsules were filled with an organic intracapsular matrix, in which 40–153 (=94) eggs were embedded. The mean diameter of the uncleaved ova was 352.5 µm. The organic intracapsular matrix and some nurse eggs were used as an extra-embryonic food source. Hatching occurred at an advanced veliger stage, immediately followed by metamorphosis. Therefore, the larval development of S. arenarius is of a lecithotrophic intracapsular type, with a short, free-swimming/crawling phase.Communicated by O. Kinne, Oldendorf/Luhe     

Early life history of two closely related gastropods,<Emphasis Type="Italic"> Rissoa auriscalpium</Emphasis> and<Emphasis Type="Italic"> Rissoa italiensis</Emphasis> (Caenogastropoda: Rissoidae)

A comparative analysis of the early life history of two common gastropod forms living on the leaves of the Mediterranean seagrass Posidonia oceanica has been carried out, taking into account timing and phases of development. The forms have a different distribution along a depth transect at Ischia Island (Gulf of Naples, Italy), the more squat and ribbed being mainly present between 1 and 15 m depth, with a maximum density at 3 m, the other, between 10 and 30 m. They were distinguished as sibling species, Rissoa italiensis (Verduin, 1985) and Rissoa auriscalpium (L., 1758), on the basis of protoconch size. However, to date, the systematic relationship between them is still unclear. Electrophoretic studies showed lower genetic distance between the two forms along the transect, than among populations from other locations in the Mediterranean Sea. R. auriscalpium have planktonic development, with a higher number of smaller eggs, while R. italiensis have an intracapsular veliger, with hatching juveniles. As for the patterns of development, while the same stages are recognizable in the two gastropod forms up to the formation of the veliger, several differences in timing occur, along with differences in growth and death rates of the embryos. R. auriscalpium hatches as a veliger at the first phase of torsion, about 10 days after spawning, while R. italiensis completes its torsion and metamorphosis within the capsule, and then hatches as a juvenile about 18 days after spawning. The time of encapsulation is what seems to lead to similar developmental sequences with different reproductive patterns and ecological roles, depending on the organization of the elements in the capsule.Communicated by R. Cattaneo-Vietti, Genova     

Embryonic development and maternal–embryo relationships of the Portuguese dogfish <Emphasis Type="Italic">Centroscymnus coelolepis</Emphasis>

The embryonic development and the level of dependency of the embryos from the maternal organism were investigated for Centroscymnus coelolepis. During the development, there is a 22–32% loss of organic matter and gains of 95–122% and 29–46% on moisture and inorganic matter, respectively. Only 28–48% of the initial yolk energy is conserved in embryos, being the remaining used in catabolism. In late pregnancy, the epithelium thickness of uterine villi decreases and vascularity increases. Results suggest that only water and minerals are transferred from the mother to the embryos. Monounsaturated fatty acids are the main type of fatty acids representing 48% of the total yolk fatty acid content, followed by polyunsaturated (30%) and saturated fatty acids (18–21%).     

Microbial activity and accumulation of organic matter in the burrow of the mud shrimp, <Emphasis Type="Italic">Upogebia major</Emphasis> (Crustacea: Thalassinidea)

Microbial activity and accumulation of organic matter in the burrow of the thalassinidean mud shrimp, Upogebia major, were studied on a tidal flat along the northern coast of Tokyo Bay, Japan. The burrow of U. major is Y-shaped with an upper U-shaped part plus a lower I-shaped part. Its lower part can extend to a depth of 2 m. In the present study, we compare electron transport system activity (ETSA), bacterial abundance and organic matter content [total organic carbon (TOC), total nitrogen (TN) and chlorophyll a (chl. a)] of the burrow wall sediment with the tidal flat surface and non-burrow sediments. We also compared the U- and I-shaped part in terms of these parameters. ETSA in the burrow wall was higher than at the tidal flat surface in the warmer season, and was always higher than at surrounding non-burrow sediments. Bacterial abundance in the burrow wall was higher than at the tidal flat surface and surrounding sediment. TOC and TN contents in the burrow wall were two to three times higher than those at the tidal flat surface and non-burrow sediments, regardless of season. However, there was no significant difference in chl. a content between burrow wall and tidal flat surface. These results suggest that organic enrichment of the burrow wall is a result of organic matter particles such as phytodebris accumulation along the burrow wall. For all parameters of the burrow walls, no statistical differences were found between the two parts. Present results indicate that U. major actively transports the water containing suspended organic particles not only through the U-part but also into the deeper I-part. These indicate that burrow of the mud shrimp provides a dynamic environment for microbial community in tidal flat sediment.     

Costs of extended brood protection in the Antarctic sea star, Neosmilaster georgianus (Echinodermata: Asteroidea)

Changes in size, morphology, and biochemical composition in adults and embryos of a brooding sea star, Neosmilaster georgianus (Sladen), were studied in a population adjacent to Anvers Island, Antarctic Peninsula (64°46′S; 64°04′W) during the austral spring, 1991. Five morphological stages of development were designated in 24 broods, and for each the weight and biochemical composition of the brooding adults and their embryos were determined. Between Stage 1 and 2, the dry weight (dw) and organic weight (ow) of the embryo did not change. From Stage 2 to 3, the dw and ow increased significantly by 10%. Stage 2 and 3 embryos were in clusters of a few (2 to 10) to as many as 40 individuals. In the smaller clusters, individual embryos were attached by tissue cords to another, sometimes atrophied, brood member. In the larger clusters, they were attached to a central mass of tissue containing remnants of embryos. We interpret these interactions as a form of cannibalism which may account for the weight gains between Stage 2 and 3. During Stages 4 and 5, as juvenile form was approached, the dw and ash weight of the young increased significantly and the ow decreased significantly. The calculated energy content for the juvenile (Stage 5) was not significantly different from the energy content of the earliest undifferentiated stage (Stage 1), an indication that most of the organic matter in the egg is the primary contribution to the large juvenile. In brooding females, pyloric caeca indices declined by 52% from Stage 1 to Stage 5 and pyloric energy stores declined by 63% due to proportionately equivalent declines in protein and lipid. The ovary index was low and increased only slightly during brood protection, while the size of the largest oocytes remained approximately 23% that of ova. Energy stores in the pyloric caeca of brooding N. georgianus thus become depleted over a long period of incubation during which the adults apparently starve. This may delay oocyte development and ultimately limit the frequency of individual reproduction. Received: 15 September 1998 / Accepted: 9 March 1999     

Developmental and genetic evidence for the existence of three morphologically cryptic species of<Emphasis Type="Italic"> Crepidula</Emphasis> in northern Chile

The taxonomy of Crepidula has been based principally on shell morphology, but shell characteristics in this genus may be strongly influenced by the substrates they inhabit and are thus of limited use in classification. Four species in this genus are currently recognized from the Chilean coast, of which two, Crepidula dilatata and C. fecunda, are sympatric. These species are morphologically cryptic and differ only in their larval development: C. dilatata has indirect development with a planktonic phase, and C. fecunda direct development within egg capsules. The other two species, C. coquimbensis and C. philippiana, are species that occur within gastropod shells inhabited by hermit crabs and differ from the other species in internal morphology of the shell, developmental features, and geographic distribution. Analyses of nine morphological characters, eight developmental characters, and 26 enzymatic loci were carried out on four Crepidula populations in the Coquimbo region of northern Chile. Four populations of Crepidula were identified based on morphological, developmental, and protein electrophoresis features. Our results suggest the presence of C. coquimbensis plus three morphologically cryptic species including C. dilatata, C. fecunda, and a new previously unrecognized cryptic species. All four populations showed differences in larval development pattern. C. fecunda have indirect development, with free-swimming larvae. The remaining three species have intracapsular metamorphosis, but different modes of larval nutrition. C. dilatata have nurse eggs that are consumed by mechanical destruction; C. coquimbensis engulf nutritional embryos; and the new species eat nutritive embryos by rotation. Protein electrophoresis data support the specific condition of each group with different larval development and endorse the use of developmental differences in taxonomy of Crepidula. Genetic distance shows that C. dilatata and C. fecunda are closely related, and that C. coquimbensis is closer to them than to the new cryptic species.     

Two modes of development in the morphospecies Crepidula dilatata (Gastropoda: Calyptraeidae) from Southern Chile

In Chinquihue Bay, a sheltered locality in Southern Chile, female Crepidula dilatata Lamarck brood egg masses which differ in intracapsular development. In some, all eggs develop and are hatched as free veliger larvae (indirect development). In others, only some of the eggs develop and hatch as young adults (direct development), the rest being consumed as nurse eggs. Two possible interpretations are considered: intrapopulation variation of developmental pattern in a species, and the coexistence of two sibling species. Stages of the intracapsular development are described and illustrated. Fecundity varies according to size of the female. For females brooding eggs with indirect development it is estimated at between 3840 and 85575 embryos per spawn; for females with eggs undergoing direct development, embryo production per egg mass is estimated at between 70 and 812. A comparison is made between females brooding the two kinds of egg masses. Minor differences exist in adult coloration, shape of egg capsules, egg diameters, spawning season and their abundance and distribution in the intertidal. Brooding females with eggs displaying indirect development are significantly larger than those with eggs of direct development. The latter females form chains subtidally but not intertidally as do females bearing eggs with indirect development. On these females with pelagic larvae, chains frequently include sedentary males which reach large sizes. On basal females with direct development, chains contain mostly other females, suggesting that matings are temporary and by errant males. These results are compared with existing information in the literature for other species both in this genus and in other marine invertebrates. It is concluded that two sympatric sibling species with different modes of development are present in the morphospecies C. dilatata Lamarck in Southern Chile.Devoted to Dr. E.F. Kilian, Zoologisches Institut der Universität Gießen, FRG.     

Influence of environmental factors and female size on reproductive output in an invasive temperate marine gastropod <Emphasis Type="Italic">Rapana</Emphasis><Emphasis Type="Italic">venosa</Emphasis> (Muricidae)

Life history and reproductive strategies influence population dynamics at the inter- and intra-specific level. Environmental conditions suitable for gonad development and spawning, the reproductive range, may be a smaller portion of the broader species distribution. The only known breeding population of veined rapa whelks (Rapana venosa) in North America is in Chesapeake Bay, USA. There is considerable interest in the potential reproductive range of this non-indigenous species given the rapa whelk’s negative impacts on commercial shellfish species in both its native and introduced ranges. Weight-specific reproductive output is described for wild caught Chesapeake Bay rapa whelks maintained in flow-through mesocosms for 2 years. Measured reproductive output within and between egg capsule deposition seasons (years) in relation to water temperature, salinity, daylength, and female size is used to describe the rapa whelk’s reproductive range. Egg capsule production is influenced by seasonal and absolute water temperatures as well as seasonal daylength cycles. Egg capsule deposition by Chesapeake Bay rapa whelks begins at water temperatures of approximately 18°C and continues for 11–15 weeks. Forty to 70% of female whelks deposited egg capsules in most weeks during this season, producing 150–200 egg capsules female⁻¹ week⁻¹. Water temperatures >28°C caused reduced egg capsule production relative to temperatures of 20–25°C. Egg capsule production was positively related to seasonal changes in daylength, and two peaks of egg capsule deposition were observed in the 2001 and 2002 deposition seasons. The combination of declining daylength and higher water temperatures in late summer was associated with the cessation of egg capsule deposition. A lower average weight specific reproductive output in 130–145 mm SL rapa whelks (average 12 ± 1%) than in 90–106 mm SL rapa whelks (average 22 ± 1% of body weight) may reflect a life history that balances the physiological costs of maintaining a large body mass with the production of many planktonic larvae from multiple clutches of egg capsules per breeding season over a 10–15-year lifespan. Estimates of the cumulative day-degree requirements corresponding to the annual initiation of egg capsule deposition were 238 and 236 for 2001 and 2002, respectively. Reproductive output and day-degree requirements for Chesapeake Bay rapa whelks were similar to values calculated from previous studies of native muricids (Eupleura caudata and Urosalpinx cinerea). A latitudinal range of 30–41° (N and S) is predicted as the realized reproductive range for rapa whelk populations on the basis of the day-degree requirements for native whelks and reproductively active invasive rapa whelk populations. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Suspended particulate organic matter in a Mediterranean submarine cave

Submarine caves display a paucity of benthic density and biomass that may be related to low trophic resources. Analysis of organic carbon, organic nitrogen, carbohydrate, protein and lipid content of suspended particulate matter was made during the period July 1985–July 1987 in a Mediterranean cave (Marseille, France) in order to determine any differences in the particulate organic matter composition along an horizontal transect. Particulate organic matter content clearly declined from the entrance of the cave to the dark inner area. This impoverishment could not be explained by a simple decrease in a few organic compounds, but appeared to be related to the combination of a decrease in both the amount and the composition of the suspended particles. Three progressive levels of impoverishment were identified towards the dark inner area of the cave: (i) decreasing amounts of seston; (ii) decreasing organic content of particles; (iii) increasing proportions of the geopolymeric (i.e., humic) components in the remaining organic matter, indicating increased degradation. The cave appeared to be sharply divided into two distinct sections — a twilight outer section whose waters contained slightly lower amounts of particulate organic matter than the open sea, and a dark inner section, 8 to 10 m higher, separated from the outer section by a steep rise and with waters of low organic matter content. The water in the twilight section appeared to be in thermal equilibrium with the open sea, and that in the dark inner section displayed thermal stratification. These differences indicated the presence of two distinct water bodies with contrasting average residence times, estimated as 1 d in the outer twilight section and 8 d in the dark inner section. The joint action of sedimentation and degradation resulted in an abrupt depletion of particulate organic matter in the dark inner section accompanied by a decrease in the benthic fauna. The decline in benthic heterotroph populations is probably related to the abrupt transition to oligotrophic conditions.     

Oogenesis and spawn-formation in the pigmy lion fish Dendrochirus brachypterus (Pteroidae)

Dendrochirus brachypterus possesses a special type of ovary in which the germinative epithelium surrounds a spongy stroma hanging in the lumen. The inner side of the ovarian wall is covered by a special type of epithelium that produces a gelatinous envelope around the gametes (spawn) during reproduction. In general, oogenesis in the lion fish resembles that of other teleosts, the characteristic difference being in the formation of ovigerous peduncles-protrusions of the spongy stroma, each of which bears a single oocyte. During oogenesis the oocytes rise above the germinal epithelium, enveloped in their follicular epithelia. The larger the oocyte the greater the length of its peduncles and the further its position from the germinal layer (Fig. 3). During this phase of development, a marked multiplication of yolk granules occurs; they increase their diameter and finally homogenize at ovulation. Under favorable thermal conditions, oogenesis continues all year round. The peduncular growth of oocytes enables the growing egg cells to move away from the egg-producing surface, thus making space available for the ovary to produce more eggs. This ability should have a survival value for these fish if we take into consideration that the most vulnerable stage in their life history is that of planktonic larvae (Fishelson, 1975).     

Carbonaceous resin capsule for vapor-phase monitoring of volatile hydrocarbons in soil: partitioning and kinetic model verification

The resin capsule system (RCS) was tested as a means of providing data on the presence and forms of volatile hydrocarbons. Results indicated that resin capsules provided data showing sensitivity to soil variables (texture and moisture content) and time. The objectives of this paper are to evaluate the RCS methodology and to determine whether carbonaceous resin capsules provide results that can be described by fundamental chemical partitioning and kinetic principles. Findings revealed a significant relationship between quantities of benzene, toluene, ethylbenzene, and xylene adsorbed on the capsule and quantities partitioned into the vapor phase. Kinetic evaluation indicated that the vapor adsorption by the resin capsule is regulated by diffusion processes. No verification of rate-limiting processes was possible due to limitations imposed by the experimental design, but it appears that during early stages, adsorption rate was limited by vapor diffusion through the soil. The resin capsule data also reflected differences that would be expected due to properties of the organic liquids present. These results provide further evidence that the RCS could be developed to suggest direct in situ monitoring to reveal quantities and nature of organic substances in soils.     

Gradients in biodiversity and macroalgal wrack decomposition rate across a macrotidal, ultradissipative sandy beach

The decomposition process of, and the meiobenthic and nematode colonization on, stranded macroalgae were studied along a macrotidal, ultradissipative sandy beach gradient at De Panne (Belgium). The horizontal patterns of macro- and meiofaunal densities and diversity in this beach have been well-documented. Defaunated Fucus detritus was buried in situ in litter bags, 10–15 cm under the sediment surface, at seven stations across the beach profile. Colonization by (meio)fauna and organic matter decomposition rates measured as carbon, nitrogen, phosphorus and dry weight loss, were studied after 10, 21 and 52 days of incubation. Algal detritus was colonized mainly by nematodes dominated by Rhabditidae, Sabatieria, Dorylaimoidea, Monhystera, Paracanthonchus and Daptonema. Organic matter weight loss was slowest at the higher and lower shore and fastest in the mid-shore, in line with the horizontal pattern of nematode diversity in the beach sediment. However, there was no consistent relationship between nematode diversity inside the litter bags and organic matter weight loss, prohibiting an unequivocal conclusion on whether the observed parallel between extant nematode diversity and organic matter decomposition rate indicates a causal relationship or follows from the fact that both are governed by the same set of physicochemical conditions. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Burrow morphology,tube formation,and microarchitecture of shell dissolution by the spionid polychaete Polydora websteri

Burrow morphology, detrital tube formation within the burrow, and ultrastructure of the surface of the burrow of Polydora websteri Hartman, 1943, are described in the shells of Crassostrea virginica (Gmelin) and of Mytilus edulis Linné. Observations were made on live worms living normally in artificial preparations of polished shell covered with transparent plastic film. P. websteri is capable of settling wherever there are crevices in shell surfaces, and slowly penetrates the shell forming a U- or flask-shaped cavity. A detrital tube is formed within the burrow from detritus collected and pressed in place by the worm. During this process, the modified setae of the 5th setiger are thrust against the wall of the inner tube and help to keep the diameter of the inner tube constant throught its length. Individuals of P. websteri burrow into the shell when placed on preparations of polished shell of Mytilus edulis under plastic film. As they do so, they secrete a viscous fluid which dissolves interprismatic and interlamellar organic matrices and then dissolves the exposed crystals. Etched prisms and lamellae reveal a complex pattern of internal dissolution. Enlargement of the burrow by the worm thus takes place by chemical dissolution of shell and also probably by flushing of loosened partially dissolved prisms and lamellae from the tube. Some shell fragments become incorporated in the walls of the detrital tube. There is no evidence of setal abrasion on the shell walls of the burrow even at the ultrastructural levelUniversity of Delaware College of Marine Studies Contribution No. 99.