Body size variation in the sexually dimorphic scaphopod Rhabdus rectius (Carpenter, 1864) (Dentaliida: Rhabdidae)

ABSTRACT

Male-biased sexual size dimorphism typically evolves via sexual selection for larger males that are favoured by choosy females or are more successful in mate competition with other males. Among marine invertebrates that broadcast their gametes into the ocean for fertilisation, this form of sexual size dimorphism is rare because such species lack direct interactions among males or between the sexes. However, the broadcast-spawning tusk shell Rhabdus rectius was recently reported to show strong male-biased sexual size dimorphism. That pattern might imply interesting and undiscovered sexual selection in this species. We found instead that the distribution of body size variation (weight, shell length) was similar between males and females of R. rectius, and mean sizes were not different between the sexes. However, we noted a male-biased sex ratio (～1:1.3) in our large sample of individuals. Many live scaphopods (and several dead shells) showed partial or complete boreholes drilled by predatory gastropods. Boreholes were observed on males and females in similar proportions. We collected scaphopods along with multiple individuals of one likely scaphopod predator, the small moon snail Euspira pallida, and in the lab we observed successful attacks by moon snails on tusk shells.     

Can Naticid Gastropod Predators Be Identified by the Holes They Drill?

We tested the hypothesis that drillholes made by different species of predatory naticid gastropods can be differentiated by variability in the inner (IBD) and outer borehole diameters (OBD) of the holes they drill. We compared two samples of Mya arenaria that were drilled by different predators, Euspira heros and Neverita duplicata, under experimental conditions. Mean IBD:OBD ratio was significantly greater for holes drilled by Euspira compared to Neverita, indicating that Euspira drills a steeper drillhole than Neverita. We also found consistent differences between the two naticids for slopes of regression lines of IBD on OBD after standardization for predator size and prey size and thickness, with slopes for Euspira being steeper, but results were not statistically significant. However, the range of IBD:OBD ratios was wide and overlapped considerably for each species, which decreases confidence in assignment of individual drillholes to a particular predator species. At least in the case of these two naticid species, interspecific differences in variation of the inner and outer diameters of the holes they drill have limited utility in identifying the maker of individual boring traces in the fossil record.     

Traces of Naticid Predation on the Gryphaeid Oyster Pycnodonte dissimilaris: Epifaunal Drilling of Prey in the Paleocene

Traces of drilling predation by naticid gastropods were observed on 51 valves of the free-lying, semi-infaunal oyster Pycnodonte dissimilaris (Gryphaeidae) from the Paleocene Hornerstown Formation, in New Jersey. Stereotypic behavior of the predator is indicated by the highly constrained placement of drill holes, 94% of which are centrally located on the oyster shells. Predator—prey mismatches in size, involving small predators that drilled through the upper valves of relatively large oysters, are documented by comparison of outer borehole diameter, as an index of predator size, with the sizes of the oyster shells. Results of this analysis suggest that at least some prey were drilled epifaunally, as they were too large to be manipulated and buried by the predator. This indicates, together with reports of epifaunal drilling by living naticids, that such behavior is geographically and stratigraphically more widespread in the Naticidae than has previously been acknowledged. This in turn suggests that epifaunal drilling of prey is a plesiomorphic, opportunistic mode of behavior, conserved in the evolution of the Naticidae, that has permitted subsequent escalation or expansion in range of naticid foraging from a more narrowly defined infaunal paradigm into exposed intertidal refugia.     

Paleobiological implications of shell repair in recent marine gastropods from the northern Gulf of California

Shell repair frequencies in eleven species of Recent gastropods from the northern Gulf of California vary with habitat, shell morphology and intensity of durophagous predation. Squat shells with large apertures tend to have high repair frequencies (0.25–0.50). Shell thickness at the aperture and shell size are not correlated with frequency of repair. Significant intraspecific variation in repair frequency exists between habitats. Samples from rocky habitats have statistically higher repair frequencies than samples of the same species from sandy habitats. However, habitat‐related variation between species is not apparent.

Trends in co‐evolution of gastropods and their durophagous predators are based on the indirect evidence of shell repair frequencies through time. Variation in repair frequency due to environmental and morphological factors may obscure predator‐related temporal trends in repair frequency.     

SHAPE DIFFERENCES AMONG BOREHOLES DRILLED BY THREE SPECIES OF NATICID GASTROPODS

Naticid gastropods leave a record of predation in the form ofcharacteristic countersunk circular boreholes. We fed bivalveprey to naticids to test whether the ratio between the innerand outer borehole diameter differed among three Recent naticidspecies: Euspira heros, E. lewisii and Neverita duplicata. Fossilshells from a Miocene-aged assemblage, containing naticids previouslyidentified as E. heros and N. duplicata, were also includedin our analyses. Ratios of the inner to outer diameters of naticidboreholes in bivalve shells showed that there were significantdifferences in borehole shape between E. heros and the two otherRecent species. We found no significant differences betweenthe ratios of E. lewisii and N. duplicata; however, a principalcomponent analysis showed that boreholes distinctly clusteraccording to species. The Miocene boreholes were also distinctfrom the Recent, indicating that the Miocene naticids may bea separate species from E. heros and N. duplicata. We have shownthat there is a species-specific component to borehole geometry.This may have important implications for palaeobiological studies,because the index of borehole functionality, previously describedas an inner to outer borehole ratio of 0.5 for all naticids,may differ between species. (Received 23 March 2004; accepted 10 January 2005)     

Microbial Diversity at Iron-Clay Interfaces after 10 Years of Interaction Inside a Deep Argillite Geological Formation (Tournemire,France)

In the context of a geological disposal of radioactive waste in clayey formations, the consequences of microbial activity are of concern regarding the corrosion of metallic components. The purpose of the present work was to characterize the microbial diversity that may have impacted corrosion processes at the interface between re-compacted argillite and steel coupons after 10 years of interaction inside the Toarcian argillite layer in Tournemire (France) under in situ conditions. Several types of steel coupons were introduced in 1999 in two boreholes (so-called CR6 and CR8) drilled in the geological formation and filled with re-compacted argillite. CR6 borehole was drilled horizontally from a century-old railway tunnel and coupons were placed in the undisturbed argillite. CR8 borehole was drilled vertically under the tunnel, in conditions influenced by the draining of the Cernon fault water. CR6 and CR8 boreholes were overcored 10 years later and steel coupons as well as re-compacted argillite samples were analyzed separately. The characterization of their microbial diversity was carried out using culture media and molecular methods using 16S rRNA genes cloning. Data resulting from both approaches were complementary. Isolates and clone sequences were affiliated to only 3 bacterial phyla: Firmicutes, Actinobacteria and Proteobacteria. The biodiversities differed depending on the steel type and the borehole considered, indicating the influence of both iron-clay interactions and in situ environmental conditions. This analysis has highlighted the presence of sulphate-reducing bacteria, iron-reducing bacteria and isolates capable to develop at high temperatures. These microorganisms can grow at the interfaces between materials in a very short period of time compared with planned durations of disposal. Thus, these results should be considered to assess the consequences of microbial activities on the evolution of the metallic components like overpacks.     

Predation by juvenile Polinices duplicatvs (Say) on Gemma gemma (Totten)

Feeding by juvenile Polinices duplicatus (Say) on Gemma gemma (Totten) at Barnstable Harbor, Massachusetts, was examined using laboratory experiments and collections of naturally occurring bored shells. Snails < 19 mm fed on Gemma in the laboratory and borehole diameter was directly related to predator size. Field collections of drilled shells showed that Gemma was an important prey of 0-year-class Polinices. Boreholes in Gemma shells from field collections were made primarily by snails of 1– mm, and < 3% were made by snails of > 10 mm. Most bored Gemma were large 1- to 2-yr-old individuals. The proportion of empty Gemma shells containing boreholes ranged from 3.7–14.6%, indicating that naticid prédation was a minor source of total Gemma mortality. Collections of bored shells closely reflected both the size range and relative abundance of natural predators of Gemma.     

Strengthening Borehole Configuration from the Retaining Roadway for Greenhouse Gas Reduction: A Case Study

A monitoring trial was carried out to investigate the effect of boreholes configuration on the stability and gas production rate. These boreholes were drilled from the retaining roadway at longwall mining panel 1111(1) of the Zhuji Coalmine, in China. A borehole camera exploration device and multiple gas parameter measuring device were adopted to monitor the stability and gas production rate. Research results show that boreholes 1~8 with low intensity and thin casing thickness were broken at the depth of 5~10 m along the casing and with a distance of 2~14 m behind the coal face, while boreholes 9~11 with a special thick-walled high-strength oil casing did not fracture during the whole extraction period. The gas extraction volume is closely related to the boreholes stability. After the stability of boreholes 9~11 being improved, the average gas flow rate increased dramatically 16-fold from 0.13 to 2.21 m³/min, and the maximum gas flow rate reached 4.9 m³/min. Strengthening boreholes configuration is demonstrated to be a good option to improve gas extraction effect. These findings can make a significant contribution to the reduction of greenhouse gas emissions from the coal mining industry.     

Ostracods from the Upper Cretaceous–Paleocene Jiaozhou Formation of Jiaozhou,Shandong, China and their biostratigraphic significance

Late Mesozoic–Cenozoic non-marine deposits are well developed in the Jiaolai Basin, eastern Shandong Province, China, yielding abundant fossils including ostracods. In order to further understand the geological settings of the basin during the Late Cretaceous–Paleogene, three boreholes (JK-1, JZK-01 and JZK-02) were drilled in the city of Jiaozhou. Nine genera and sixteen species (including six indeterminate species) of ostracods are described from the Jiaozhou Formation in the three boreholes, including Cypridea, Mongolocypris, Talicypridea, Candona, Eucypris, Cyprois, Porpocypris, Sinocypris and Timiriasevia. Of these, Porpocypris sphaeroidalis Guan, 1978 is reported from northern China for the first time, indicating that this species has great potential to be an indicator of the K/Pg boundary. Correlation of these species with the known Cretaceous–Paleogene ostracod assemblages suggests that the age of the Jiaozhou Formation could be latest Cretaceous–Paleocene in borehole JK-1, Campanian–Maastrichtian in borehole JZK-01, and Maastrichtian–Danian in borehole JZK-02. The exact position of K/Pg boundary, however, is unavailable for the present in the Jiaozhou Formation from boreholes JK-1 and JZK-01.     

Stereotypic and size-selective predation in Polinices pulchellus (Gastropoda: Naticidae) Risso 1826

Polinices pulchellus were size-selective in their choice of Cerastoderma edule. Large predators (12-15.9 mm shell length) selected both larger and a wider size range of cockles than smaller individuals (4-11.9 mm shell length). Considerable overlap occurred in the sizes of cockles frequently drilled by different size classes of snails, indicating that certain sizes of cockles may be most profitable to a wide range of predator sizes. Consumption rates were highest during July and August and were closely related to seawater temperature. Inner and outer drill hole diameters were both correlated with predator size, and the morphology of the drill hole was geometrically similar across a range of predator sizes. Polinices pulchellus showed no preference for either the left or right valve and drilled most cockles in the centre of the shell valve. The relationship between the distance of the drill hole from the umbo and prey size was unaffected by predator size, such that predators of different sizes were not found to drill cockles in different positions. When disturbed during drilling, incomplete drill holes were abandoned and, when drilling resumed, it occurred in new locations on the surface of the shell valve. The findings of this study highlight the stereotyped nature of drilling behaviour seen in the family Naticidae.     

Investigation of the potential for microbial contamination of deep granitic aquifers during drilling using 16S rRNA gene sequencing and culturing methods

Total number of bacteria, viable counts of aerobic and anaerobic heterotrophic bacteria and 16S rRNA gene diversity were investigated during drilling of three boreholes in the walls of the Äspö hard rock laboratory tunnel, at depths ranging from 380 to 446 m below sea level. Water samples were taken from the drill water source, the drilling equipment and from the drilled boreholes. The drill water was kept under nitrogen atmosphere and all equipment was steam cleaned before the start of a new drilling. Total and viable counts of bacteria in the drilled boreholes were several orders of magnitude lower than in the samples from the drilling equipment, except for sulphate reducing bacteria. A total of 158 16S rRNA genes that were cloned from the drill water source, the drilling equipment and the drilled boreholes were partially sequenced. The drilled boreholes generally had a 16S rRNA diversity that differed from what was found in samples from the drilling equipment. Several of the sequences obtained could be identified on genus level as one of the genera Acinetobacter, Methylophilus, Pseudomonas and Shewanella. In conclusion, the tubing used for drill water supply constituted a source of bacterial contamination to the rest of the drilling equipment and the boreholes. The results show, using molecular and culturing methods, that although large numbers of contaminating bacteria were introduced to the boreholes during drilling, they did not establish in the borehole groundwater at detectable levels.     

Predatory boreholes in Tournaisian (Lower Carboniferous) spiriferid brachiopods

A brachiopod fauna from the uppermost part of the Tournaisian Tournai Formation (Belgium) contains an undetermined species of Crurithyris (Spiriferida, Ambocoeliidae), which displays numerous bored shells. About 8% of the 432 specimens with conjoined valves display single, small (≤ 1 mm) boreholes, which are smooth-sided, cylindrical or weakly conical, circular to slightly elliptical in plan view, perpendicular to the shell surface and generally complete. Of the 35 bored articulated specimens, 27 were drilled on the ventral valve. Most of the boreholes are located in the posterior half of the shell, and no case of edge-drilling has been observed. The boreholes were drilled by a predator, or possibly a parasite, which selected individuals greater than 2.5 mm long. Crurithyris sp. may have represented an attractive (in terms of energy cost) and easy target for a small-sized predator because of its thin shell and ornament of minute spines.     

Drilling under threat: An experimental assessment of the drilling behavior of Nucella lamellosa in the presence of a predator

Using the drilling muricid, Nucella lamellosa (Gmelin 1791) and its prey, the mussel Mytilus trossulus (Gould 1850), the impact of a secondary predator, the crab Cancer gracilis, on drilling was investigated experimentally. The frequency of incomplete holes was compared under two conditions: (1) when the gastropod's natural predator, C. gracilis, was present and (2) when it was absent. The results indicate that the presence of a secondary predator can affect drilling activity, leading to a significant increase in the frequency of incomplete drill holes. The introduction of a secondary predator can also decrease the overall drilling frequency. The size distributions of completely and incompletely drilled mussels suggest that in the presence of the secondary predator the decision by the gastropod to either abandon or continue drilling its prey might be influenced by how much time it has already invested into drilling or the size of the prey item. These results are important for the ecological and evolutionary implications of incomplete drill holes frequencies, especially with regard for their use as proxies for evolutionary prey improvement.     

Predation on freshwater snails in Miocene Lake Steinheim: a trigger for intralacustrine evolution?

Shells of the freshwater gastropods Gyraulus trochiformis (Planorbidae or ramshorn snails) and Bania pseudoglobula (formerly Pseudamnicola, Hydrobiidae) from Miocene Steinheim Basin in SW Germany contain small holes with a mean diameter of 0.8 mm. Analyses of comparable holes are so far unknown from fossil or extant assemblages of freshwater shells. This analysis of the perforated shells suggests that the holes were not formed by post‐depositional or pure taphonomic processes. Instead, they were most likely produced by predators. This analysis widens the means for identification of predation on freshwater snail shells that can be used in other palaeolimnological studies. The co‐occurrence of fish teeth and perforated shells in the studied samples as well as the perforation features suggest that the predator was either barbel or tench fish. The correlation between shell sizes and hole diameters suggests a size relationship between predator and prey that may generally be related to gape‐limited fish predators. The co‐occurrence of perforated shells in these lake sediments with a dominance of large and armoured shells suggest that these larger shells with protuberances and knobs could not be crushed by the gape‐limited fish predators. This analysis is the basis for a hypothesis that the endemic evolution of Gyraulus in Lake Steinheim, with some varied forms of shell thickness and morphology, was triggered by a predator–prey relationship based on adaptations to avoid shell‐breaking predators.     

Probable predatory borings in late Cretaceous bryozoans

Borings are described in zooids of Cenomanian to Campanian melicerititids, an aberrant group of cyclostome bryozoans with calcified opercula. The borings are circular or elliptical, straight-sided and have a diameter of 40–90 μn. Autozooids were drilled in preference to heterozooids and most borings penetrate the operculum. The shape and distribution of boreholes suggests that they were made by a predator attacking one zooid at a time. The predator responsible may have been a gastropod, most probably a nudibranch or a micromorphic muricid. Bryozoa, Gastropoda, predation, borings, Upper Cretaceous.     

Macroecology and evolution of a crab ‘super predator’, Menippe mercenaria (Menippidae), and its gastropod prey

It is widely accepted that the fossil record shows both the evolution of more powerful durophagous marine predators through time and, in response, major shifts in life mode and morphology for many prey taxa. Few fossil studies, however, have successfully identified particular predator species with respect to causing evolutionary change in particular prey species. We present evidence that the evolutionary appearance in the western Atlantic of the stone crab, Menippe mercenaria, an extraordinarily powerful durophagous predator, contributed to the appearance of sinistrality, which is very rare, in two genera of marine gastropods (Conus and Sinistrofulgur) during the Pliocene. Based on this conclusion, we suggest that modern fishing pressure on stone crabs may lead to evolutionary changes in their present day prey.     

Drilling predation on scaphopods and other molluscs from the Upper Cretaceous of Manitoba, Canada

Abundant information on drilling predation upon fossil bivalves, gastropods, and brachiopods has been accumulated during the past several decades, but very little is known about the predation on marine, infaunal carnivorous scaphopods. A survey of over 440 specimens of the scaphopod Dentalium gracile collected from the Upper Cretaceous Millwood Member (Campanian) of the Pierre Shale at a site near Russell, Manitoba displays a drilling percentage of approximately 35%. This figure is higher than previously reported for the scaphopods of similar age elsewhere, but is comparable to or lower than that of the sub-Holocene (34–61%). The difference in drilling percentage among different collections may be taxon-related or affected by the composition and structure of the fossil community. Morphologically, the drill-holes, produced by predatory drilling, are beveled with a countersunk profile of clean sharp edges. The drill-hole inner margin is ovate whereas the outer margin is nearly circular. Among the Manitoba specimens, drill-holes seem to be more common on the lateral side. Presumably due to the lack of effective defense strategy, the prey effectiveness is low (∼3.2%). There is no correlation between drill-hole dimension and scaphopod prey size, indicating that predator size does not correlate with the prey size. Nearly 480 specimens of cephalopods, pelecypods, and gastropods were also collected from the same community. There were only a total of 16 drill-holes on this additional material. Over 400 specimens of the only naticid Euspira obliquata were recovered from the same site and are believed to be the predators of the scaphopods and other molluscs. The low percentage (∼2.5%) of drilling on the abundant gastropod E. obliquata may indicate mistaken or opportunistic attacks by the predator, or that the cannibalism is not common in this community.     

Effects of vertebrate predation on freshwater gastropods: an exclosure experiment

A field experiment with predator exclosures was performed in a pond in Southern Sweden to evaluate the effects of vertebrate predators on the diversity, density and distribution of freshwater gastropods. Ten exclosures (1 × 1 × 1.5 m) were erected in the nearshore vegetation at a waterdepth of 0.5–0.7 m in early June 1983 so that they included different amounts and species of macrophytes. The exclosures and cageless controls were sampled for macrophytes and gastropods at the end of the summer. The density of gastropods in the controls, exposed to predators, was positively correlated to the abundance of macrophytes. No such correlation existed in the exclosures. Further, the density of gastropods was significantly higher in the exclosures than predicted by the regression between gastropods and macrophytes in the controls. There was no difference between species composition of gastropods between controls and exclosures, but the number of species and the density of gastropods were lower in the macrophyte stands dominated byScirpus lacustris than in the more complex stands dominated bySparganium erectum. These results indicate that vertebrate predation is a major structuring force of benthic freshwater gastropod communities.     

Comparing predatory drillholes to taphonomic damage from simulated wave action on a modern gastropod

Marine drilling predation, in which the predator bores a hole through shelled invertebrate prey, plays a role in the structure of benthic communities. As drilling often leaves the prey shell otherwise undamaged, the resulting holes are also an excellent proxy for drilling predation pressure in the fossil record. Considering that a large number of predation studies focus on drilling predation in the fossil record, it is crucial that we are able to distinguish true drilling predation from taphonomy. The purpose of this study is to determine damage on Olivella biplicata shells, drilled by naticid gastropods, is distinguishable from taphonomically produced damage to these shells. In addition, the potential for preferential breakage due to either the presence or whether absence of a drillhole was investigated. Drilled and non-drilled O. biplicata shells were tumbled to simulate wave action and were checked at intervals to record accumulated damage. Drilled and non-drilled shells do not show a significant difference in damage accumulated while undergoing simulated wave action. Taphonomic damage is unlikely to be mistaken for drilling damage, due to the jagged, irregular appearance of taphonomically produced holes.