Broken pieces: can variable ecological interactions be deduced from the remains of crab attacks on bivalve shells?

Shell fragmentation patterns that result from attacks by durophagous predators on hard‐shelled marine invertebrates are a rich source of indirect evidence that have proved useful in interpreting predation pressure in the fossil record and recent ecology. The behaviour and effectiveness of predators are known to be variable with respect to prey size. It is less well understood if variable predator–prey interactions are reflected in shell fragmentation patterns. Therefore, we conducted experimental trials to test the behavioural response of a living crab, Carcinus maenas, during successful predatory attacks on the blue mussel Mytilus edulis on two prey size categories. Further, we examined resultant shell fragments to determine whether specific attack behaviours by C. maenas could be successfully deduced from remaining mussel shells. In contrast to previous studies, we observed no significant differences in attack behaviour by the predators attributable to prey size. In most experimental predation events, crabs employed an ad hoc combination of five mechanisms of predation previously described for this species. We identified seven categories of shell breakage in predated mussels, but none of these were unambiguously correlated with specific attack behaviour. Combined attack behaviours may produce shell breakage patterns that have previously been assumed to be attributable to a single behaviour. While specific patterns of shell breakage are clearly attributable to durophagy, the results of this study provide important insights into the limitations of indirect evidence to interpret ecological interactions.     

Early Cambrian record of failed durophagy and shell repair in an epibenthic mollusc

Predation is arguably one of the main driving forces of early metazoan evolution, yet the fossil record of predation during the Ediacaran-Early Cambrian transition is relatively poor. Here, we present direct evidence of failed durophagous (shell-breaking) predation and subsequent shell repair in the Early Cambrian (Botoman) epibenthic mollusc Marocella from the Mernmerna Formation and Oraparinna Shale in the Flinders Ranges, South Australia. This record pushes back the first appearance of durophagy on molluscs by approximately 40Myr.     

Shell composition of Terreneuvian tubular fossils from north‐east Sichuan,China

Identification of the primary constituents of small shelly fossil (SSF) shells is important for explaining the evolution of SSF faunas. The characteristics and constituents of Terreneuvian tubular SSFs found in north‐east Sichuan, China, are revealed using light microscopy, scanning electron microscopy and energy dispersive X‐ray spectroscopy. Petrographic thin sections indicate that the chemical composition of the shells is mainly calcium carbonate with smaller amounts of phosphorus, silica and pyrite. Most of the tubular shells composed of calcium carbonate have a distinct layered structure. Evidence of replacement of the original shell by phosphatization, pyritization and silicification, and recrystallization of calcium carbonate have been found, all of which destroyed the shell's original layered structure. Most fossils treated with acetic acid are phosphatic casts or steinkerns, with some preserving organic textures of the shell as phosphatic casts. We conclude that the Terreneuvian tubular SSFs from north‐east Sichuan were originally composed mainly of calcium carbonate; indeed, most Terreneuvian small skeletal fossils appear to have had an originally calcareous composition. The fossil casts or internal core fossils are composed of phosphate, which is related to local taphonomic processes.     

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.     

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.     

Small carbonaceous fossils (SCFs) from the Terreneuvian (lower Cambrian) of Baltica

We describe a new assemblage of small carbonaceous fossils (SCFs) from diagenetically minimally altered clays and siltstones of Terreneuvian age from the Lontova and Voosi formations of Estonia, Lithuania and Russia. This is the first detailed account of an SCF assemblage from the Terreneuvian and includes a number of previously undocumented Cambrian organisms. Recognizably bilaterian‐derived SCFs include abundant protoconodonts (total‐group Chaetognatha), and distinctive cuticular spines of scalidophoran worms. Alongside these metazoan remains are a range of protistan‐grade fossils, including Retiranus balticus gen. et sp. nov., a distinctive funnel‐shaped or sheet‐like problematicum characterized by terminal or marginal vesicles, and Lontohystrichosphaera grandis gen. et sp. nov., a large (100–550 μm) ornamented vesicular microfossil. Together these data offer a fundamentally enriched view of Terreneuvian life in the epicratonic seas of Baltica, from an episode where records of non‐biomineralized life are currently sparse. Even so, the recovered assemblages contain a lower diversity of metazoans than SCF biotas from younger (Stage 4) Baltic successions that represent broadly equivalent environments, echoing the diversification signal recorded in the coeval shelly and trace‐fossil records. Close comparison to the biostratigraphical signal from Fortunian small shelly fossils supports a late Fortunian age for most of the Lontova/Voosi succession, rather than a younger (wholly Stage 2) range.     

Do predatory drill holes influence the transport and deposition of gastropod shells?

Predatory gastropod drill holes are an abundant and easily identifiable signal of predation in ancient and modern molluscan shell assemblages. Many workers have used drill holes to interpret patterns of predation intensity and success in fossil assemblages. These studies are predicated on the assumption that the relative abundances of drilled and undrilled shells in an assemblage accurately reflect those of the community from which the shells were originally derived. The underlying assumption is that drilled and undrilled shells are transported into shell accumulations in the same manner. If this assumption is false, shell accumulations do not represent taphonomically unbiased samples, but rather preferentially sorted deposits from which conclusions about drilling predation cannot be made. To test the hypothesis that drilled and undrilled gastropod shells transport at different flow velocities, multiple transport trials were conducted on two morphologically distinct taxa, Olivella biplicata and Euspira lewisii. Individual specimens were placed in a recirculating flume tank and observed from rest (in stable orientation) until they were transported downstream. During each trial, flow velocity was slowly and incrementally increased, so as to avoid pulses of acceleration, until shells began to move downstream. Drilled and undrilled specimens of both taxa demonstrate statistically significant correlations between shell mass and average transport velocity. Similarly sized drilled and undrilled specimens of both taxa do not exhibit significant differences in transport velocity. These results indicate that predatory drill holes do not change the hydrodynamic properties of gastropod shells. Therefore, gastropod shell assemblages are not likely to be affected by differential transport and sorting of drilled and undrilled shells.     

The gnathobasic spine microstructure of recent and Silurian chelicerates and the Cambrian artiopodan Sidneyia: Functional and evolutionary implications

Gnathobasic spines are located on the protopodal segments of the appendages of various euarthropod taxa, notably chelicerates. Although they are used to crush shells and masticate soft food items, the microstructure of these spines are relatively poorly known in both extant and extinct forms. Here we compare the gnathobasic spine microstructures of the Silurian eurypterid Eurypterus tetragonophthalmus from Estonia and the Cambrian artiopodan Sidneyiainexpectans from Canada with those of the Recent xiphosuran chelicerate Limulus polyphemus to infer potential variations in functional morphology through time. The thickened fibrous exocuticle in L. polyphemus spine tips enables effective prey mastication and shell crushing, while also reducing pressure on nerve endings that fill the spine cavities. The spine cuticle of E. tetragonophthalmus has a laminate structure and lacks the fibrous layers seen in L. polyphemus spines, suggesting that E. tetragonophthalmus may not have been capable of crushing thick shells, but a durophagous habit cannot be precluded. Conversely, the cuticle of S. inexpectans spines has a similar fibrous microstructure to L. polyphemus, suggesting that S. inexpectans was a competent shell crusher. This conclusion is consistent with specimens showing preserved gut contents containing various shelly fragments. The shape and arrangement of the gnathobasic spines is similar for both L. polyphemus and S. inexpectans, with stouter spines in the posterior cephalothoracic or trunk appendages, respectively. This differentiation indicates that crushing occurs posteriorly, while the gnathobases on anterior appendages continue mastication and push food towards and into the mouth. The results of recent phylogenetic analyses that considered both modern and fossil euarthropod clades show that xiphosurans and eurypterids are united as crown-group euchelicerates, with S. inexpectans placed within more basal artiopodan clades. These relationships suggest that gnathobases with thickened fibrous exocuticle, if not homoplasious, may be plesiomorphic for chelicerates and deeper relatives within Arachnomorpha. This study shows that the gnathobasic spine microstructure best adapted for durophagy has remained remarkably constant since the Cambrian.     

Experimental tumbling of <Emphasis Type="Italic">Dreissena polymorpha</Emphasis>: implications for recognizing durophagous predation in the fossil record

Shell damage left by predators constitutes an important source of information on predator–prey interactions. However, recognition of the origins of shell damage can often be controversial, and needs to be assessed cautiously. More specifically, differentiation between predation- and abiotic-induced shell damage remains challenging. Here, we show the results of tumbling experiments using a bivalve species Dreissena polymorpha in order to determine rates and patterns of shell damage induced by physical forces in high-energy conditions. It is demonstrated that, in contrast to durophagous fish and crab predation, abiotic-induced fragmentation and damage are typically characterized by the presence of distinct abrasive scratches and wear scars on the surface of shell fragments. Furthermore, fragmented shells typically reveal a wide size distribution, and a different degree of sphericity and roundness resulting from abrasion. Importantly, large shell fragments commonly display smooth edges. These data suggest that durophagous predation, which typically induces fragmentation into large and angular shell fragments bearing no wear scars, can be reliably recognized both in present-day environments and in the fossil record.     

Invasion of a naticid predator and associated changes in death assemblages of bivalve prey in northern Japan: implications for palaeoecological studies

The recent invasion of a naticid predator (Laguncula pulchella) and associated changes in the death assemblages of bivalve prey (Ruditapes philippinarum) provide a baseline for interpreting predator–prey interactions in the fossil record. This article presents quantitative data on size‐frequency distributions (SFDs) of living and death assemblages, prey size selectivity and drillhole site selectivity from the Tona Coast, northern Japan. Before the appearance of the predator, the SFD of the death assemblage exhibited a right‐skewed platykurtic distribution, and there were very few predatory drillholes. Once the predator appeared, frequencies of predatory drillholes increased, particularly in the smallest size class (2–10 mm shell length). Furthermore, juvenile peaks in the SFDs of death assemblages sharpened, and thus, SFDs exhibited strongly right‐skewed leptokurtic distributions. These changes suggest that intense naticid predation precluded juvenile clams from growing to adulthood, and thus, many dead shells of juvenile clams were introduced into the sediment. The changes in SFDs may also indicate intensification of predation pressure in the fossil record. No temporal shifts in prey size selectivity and drillhole site selectivity were noted, despite substantial changes in the demographics of Ruditapes philippinarum. This suggests that lack of specific size classes of preferred prey species is unlikely to be a primary factor accounting for size mismatches between predator and prey, because, in such situations, naticid predators probably attack other prey species. Therefore, such a factor is unlikely to primarily explain the less stereotypical predatory behaviour (i.e. low prey size selectivity and low drillhole site selectivity), which has been frequently recognized in fossil assemblages. Such less stereotypical predatory behaviour in fossil assemblages is likely to be explained by other factors, such as the existence of multiple predator taxa and lack of specific size classes of all available prey.     

Periodic shell decollation as an ecology-driven strategy in the early Cambrian Cupitheca

Shell decollation is a growth strategy that has been adopted by a number of invertebrate taxa to offset the metabolic and ecological disadvantages of shell growth. However, little is known about the origin and evolution of this process. We here describe well-preserved specimens of the hyolith Cupitheca decollata sp. nov. preserving the decollation process, from the early Cambrian Yu'anshan Formation (c. 518 Ma) of South China. Based on a large number of specimens collectively representing different developmental stages, we use high-resolution X-ray microtomography and scanning electronic microscopy to reconstruct the process of decollation in this taxon. Cupitheca is among the earliest known small shelly fossils, and thus our discovery confirms that periodic decollation had evolved by the onset of the Cambrian explosion, reflecting the high intensity of the predator–prey arms race in early Cambrian ecosystems. A comparison between the decollation processes of Cupitheca and other shelly invertebrates suggests that periodic decollation and the associated molecular mechanisms of calcium dissolution, uptake, allocation and deposition may have had multiple independent origins.     

DUROPHAGOUS PREDATION ON MIDDLE JURASSIC MOLLUSCS, AS EVIDENCED FROM SHELL FRAGMENTATION

Abstract:  Durophagous (shell-crushing) predation is known from the beginning of the Phanerozoic, but it has been suggested that modern intensity was not reached until the Late Cretaceous and Early Cenozoic, when specialized marine durophagous taxa increased in diversity. In this paper, evidence of durophagous predation on Middle Jurassic communities of molluscan prey is presented on the basis of distinct accumulations of fossil remains in the Polish Jura (south-central Poland) that contain characteristic, angular shell fragments with sharp, non-abraded margins. The diverse fossil content of the accumulations studied, consisting of either benthic or nektic/nekto-benthic taxa, indicates that the potential predatory taxon was an opportunistic generalist, most probably fish. On the basis of taphonomic observations, the faunal accumulations are interpreted to represent regurgitated remains (pellets). The common occurrence of such accumulations in the Middle Jurassic clays of the Polish Jura indicates that durophagous predation has been intense since the mid-Mesozoic, at least locally.     

EVIDENCE FROM THE FOSSIL RECORD OF AN ANTIPREDATORY EXAPTATION: CONCHIOLIN LAYERS IN CORBULID BIVALVES

Conchiolin layers, organic-rich laminae, are characteristic of the shells of corbulid bivalves. The retention of these layers, despite their high metabolic cost, throughout the evolutionary history of Corbulidae has prompted the proposal of several adaptive scenarios to explain the origin and maintenance of these layers. The most widely held hypothesis contends that conchiolin layers are an adaptation for inhibiting drilling by predatory naticid gastropods. However, others suggest that the layers are adaptations to retard shell dissolution in waters undersaturated with calcium carbonate or to increase shell strength in the face of durophagous (shell crushing) predators. In this paper, experiments using recent Corbula (Varicorbula) gibba (Olivi) and observations of corbulids' present natural habitat demonstrate the current utility of conchiolin layers for all three functions: retardation of shell dissolution in waters undersaturated in calcium carbonate, increase of mechanical shell strength, and inhibition of drilling by predatory naticid gastropods. Earlier analyses of the extensive history of naticid predator-corbulid prey interactions suggested that conchiolin layers were an adaptation, a feature that promotes fitness and was built by selection for its current role, for deterring naticid predators. Not only are naticid drillholes widespread in fossil and recent corbulid shells, but an unusually large number of incomplete drillholes terminate unsuccessfully at conchiolin layers. In addition, a phylogenetic analysis of the origin of conchiolin layers and its function to deter naticid predators is consistent with a hypothesis of adaptation for this function. However, this hypothesis is rejected by an examination of fossil Jurassic Corbulomima. These oldest corbulids contained conchiolin layers before the evolution of naticid drilling during the Early Cretaceous. Therefore, conchiolin layers appear to be an exaptation, characters evolved for other usages and later “coopted” for their current role, for defense against drilling predators. The layers may in fact be an adaptation to resist durophagous predation.     

Drill Holes and Predation Traces versus Abrasion-Induced Artifacts Revealed by Tumbling Experiments

Drill holes made by predators in prey shells are widely considered to be the most unambiguous bodies of evidence of predator-prey interactions in the fossil record. However, recognition of traces of predatory origin from those formed by abiotic factors still waits for a rigorous evaluation as a prerequisite to ascertain predation intensity through geologic time and to test macroevolutionary patterns. New experimental data from tumbling various extant shells demonstrate that abrasion may leave holes strongly resembling the traces produced by drilling predators. They typically represent singular, circular to oval penetrations perpendicular to the shell surface. These data provide an alternative explanation to the drilling predation hypothesis for the origin of holes recorded in fossil shells. Although various non-morphological criteria (evaluation of holes for non-random distribution) and morphometric studies (quantification of the drill hole shape) have been employed to separate biological from abiotic traces, these are probably insufficient to exclude abrasion artifacts, consequently leading to overestimate predation intensity. As a result, from now on, we must adopt more rigorous criteria to appropriately distinguish abrasion artifacts from drill holes, such as microstructural identification of micro-rasping traces.     

Coevolution of a marine gastropod predator and its dangerous bivalve prey

The fossil record of the interaction between the predatory whelk Sinistrofulgur and its dangerous hard‐shelled bivalve prey Mercenaria in the Plio‐Pleistocene of Florida was examined to evaluate the hypothesis that coevolution was a major driving force shaping the species interaction. Whelks use their shell lip to chip open the shell of their prey, often resulting in breakage to their own shells, as well as to their prey. Mercenaria evolved a larger shell in response to an intensifying level of whelk predation. Reciprocally, an increase in attack success (ratio of successful to unsuccessful attacks) and degree of stereotypy of attack position by the predator suggest reciprocal adaptation by Sinistrofulgur to increase efficiency in exploiting hard‐shelled prey. A decrease in prey effectiveness (ratio of unsuccessful to total whelk predation attempts) and an increase in the minimum boundary of a size refuge from whelk predation for Mercenaria may indicate that predator adaptation has outpaced prey antipredatory adaptation. Evolutionary size increase in Sinistrofulgur most likely occurred in response to prey adaptation to decrease the likelihood of feeding‐induced shell breakage and unsuccessful predation when encounters with damage‐inducing prey occur, coupled with (or reinforced by) an evolutionary response to the whelk's own predators. Predator adaptation to Mercenaria best explains temporal changes in whelk behaviour to decrease performance loss (shell breakage) associated with feeding on hard‐shelled prey; this behavioural change limits attacks on prey to when the whelk's shell lip is thickest and most resistant to breakage. Despite evidence of reciprocal adaptation between predator and prey, the contribution of Mercenaria to Sinistrofulgur evolution is likely only a component of the predator's response to dangerous bivalve prey. This study highlights the importance of understanding the interactions among several species in order to provide the appropriate context to test evolutionary hypotheses about any specific pair of species. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 80 , 409–436.     

Killing them softly: Ontogeny of jaw mechanics and stiffness in mollusk-feeding freshwater stingrays

Durophagous predators consume hard-shelled prey such as bivalves, gastropods, and large crustaceans, typically by crushing the mineralized exoskeleton. This is costly from the point of view of the bite forces involved, handling times, and the stresses inflicted on the predator's skeleton. It is not uncommon for durophagous taxa to display an ontogenetic shift from softer to harder prey items, implying that it is relatively difficult for smaller animals to consume shelled prey. Batoid fishes (rays, skates, sawfishes, and guitarfishes) have independently evolved durophagy multiple times, despite the challenges associated with crushing prey harder than their own cartilaginous skeleton. Potamotrygon leopoldi is a durophagous freshwater ray endemic to the Xingu River in Brazil, with a jaw morphology superficially similar to its distant durophagous marine relatives, eagle rays (e.g., Aetomylaeus, Aetobatus). We used second moment of area as a proxy for the ability to resist bending and analyzed the arrangement of the mineralized skeleton of the jaw of P. leopoldi over ontogeny using data from computed tomography (CT) scans. The jaws of P. leopoldi do not resist bending nearly as well as other durophagous elasmobranchs, and the jaws are stiffest nearest the joints rather than beneath the dentition. While second moment has similar material distribution over ontogeny, mineralization of the jaws under the teeth increases with age. Neonate rays have low jaw stiffness and poor mineralization, suggesting that P. leopoldi may not feed on hard-shelled prey early in life. These differences in the shape, stiffness and mineralization of the jaws of P. leopoldi compared to its durophagous relatives show there are several solutions to the problem of crushing shelled prey with a compliant skeleton.     

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.     

MORPHOLOGY AND BEHAVIOR OF CRABS AND GASTROPODS FROM LAKE TANGANYIKA,AFRICA: IMPLICATIONS FOR LACUSTRINE PREDATOR-PREY COEVOLUTION

The shells of most lacustrine gastropods are typically small, weakly calcified, and modestly ornamented to unornamented. Similarly, most lacustrine crabs are usually small detritivores with weak chelae. A number of invertebrate taxa in Lake Tanganyika, however, deviate from these generalities. This study explores a predator-prey coevolution model as an explanation for the large, heavily calcified, and ornate gastropods and the robust, durophagous crabs of Lake Tanganyika. The endemic thiarid and viviparid gastropods from Lake Tanganyika have significantly thicker shells and higher frequencies of terminal apertural lip thickening than closely related cosmopolitan taxa from outside the lake. Tanganyikan gastropods also display considerably higher incidence of shell repair, following nonlethal shell damage, than cosmopolitan taxa of the same families. There is a strong positive correlation between gastropod apertural lip thickness and shell repair frequency among all the gastropod species analyzed. The endemic Tanganyikan potamonautid crab Platytelphusa armata (a molluscivore) possesses larger, more robust crushing chelae than other African potamonautid or potamonid crabs. In contrast with the cosmopolitan African crabs, the Tanganyikan crabs display molariform, rather than serrate dentition on their crushing chelipeds. In shell-crushing experiments, the Tanganyikan gastropod shells were an order of magnitude stronger than typical lacustrine gastropod shells, many well within the range of tropical marine gastropod shell strengths. Predation experiments with the endemic gastropods Spekia, Neothauma, Lavigeria spp., Paramelania spp. and the crab Platytelphusa armata showed that increased size, apertural lip thickness or shell sculpture reduced the successful predation rate of P. armata. Crabs with large chelae have a greater ratio of successful: unsuccessful attacks than crabs with small chelae. Among cases of successful predation, crabs with large chelae employed predation methods that required less time and energy (such as crushing the shell in the cheliped) than the methods employed by crabs with small chelae (such as peeling the shell from the aperture or the spire). The morphological, shell-crushing, and aquarium experiment data, considered in concert, provide strong support for the idea that the endemic gastropods and crabs of Lake Tanganyika have coevolved over the past 7 million years.     

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.