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.     

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.     

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.     

Additive interactions between the moon snail Euspira heros and the sea star Asterias forbesi, two predators of the surfclam Spisula solidissima

A laboratory experiment was conducted to determine whether the sea star Asterias forbesi and the naticid gastropod Euspira heros feed on surfclams, Spisula solidissima, in an additive or non-additive manner. Predators were allowed to feed on clams with conspecifics and in the presence of the other predator species. Clam mortality (measured as the rate of decline of clam number) and predator feeding rates were noted. To determine the effects of temperature on interactions among the predators, the experiment was conducted at three different temperatures. At all temperatures, feeding rate of each predator was not affected by the presence of the other species, and clam mortality in the presence of both predators was predictable from mortality in the presence of a single predator species. These additive interactions are most likely a result of habitat partitioning between the predators, with naticid snails being infaunal and sea stars being epifaunal. Previous studies in a variety of systems show no clear pattern of occurrence of non-additive interactions. Relatively small differences in predator or prey behavior may be responsible for the presence or absence of non-additive interactions. Received: 6 August 1998 / Accepted: 25 January 1999     

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.     

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.     

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.     

Pulling or drilling, does size or species matter? An experimental study of prey handling in Octopus dierythraeus ()

The influence of both predator and prey size on the shift from a pulling to a drilling predatory response was examined in the intertidal octopus Octopus dierythraeus, using an experimental program. Additionally, selective drilling, where particular regions of the prey are targeted, was examined for a variety of bivalve and gastropod prey. O. dierythraeus always initially attempted to pull bivalves apart. Shells that were eventually drilled were always subjected to significantly more pulling attempts than those that could be pulled apart, indicating that octopus are willing to expend more energy to access the flesh quickly. There was no defined threshold where bivalve size caused an octopus to switch from a pulling to a drilling response. Instead, there was a broad size range where the octopus could adopt either handling method and it varied for each individual. Octopus may only able to pull open bivalves before the molecular ratchet or ‘catch’ mechanism that many bivalves possess is engaged. This might explain the lack of a relationship between either octopus or bivalve size and the success of pulling, as it is likely that when the bivalves were presented to individual octopus they were either setting the ‘catch’ mechanism, or had already engaged it. O. dierythraeus demonstrated selective drilling on a variety of molluscan prey, with penetration sites differing between prey species. O. dierythraeus targeted the valve periphery, which was the thinnest part of the shell, therefore minimizing handling time. O. dierythraeus always drilled gastropods, but did not target the thinnest regions of the shells, with drill site varying according to the morphology of the prey. Elongate species with pronounced aperture lips were drilled in the apical region, close to the columella on the side of the opercula whereas nonelongate species were drilled immediately above the aperture. The location of drilling sites may represent a trade-off between targeting the most effective places to inject paralyzing secretions and the mechanically simplest places to drill.     

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.     

Invasion of Laguncula pulchella (Gastropoda: Naticidae) and predator–prey interactions with bivalves on the Tona coast, Miyagi prefecture, northern Japan

On the Tona coast, Miyagi prefecture, northern Japan, interactions between the alien predator Laguncula pulchella and its bivalve prey were explored using annually collected quadrat samples over a 10 year period, from 2001 to 2010. A single L. pulchella individual was first recorded in 2002, and the density increased 12-fold from 2002 to 2004. In contrast, population densities of Ruditapes philippinarum and Macoma incongrua rapidly decreased during this interval. Based on frequency of predatory drill holes on the dead shells, more than 35 % of Ruditapes philippinarum and 20 % of Macoma incongrua died because of naticid predation after 2004, while Pillucina pisidium was less vulnerable to naticid predation. L. pulchella focused attacks on P. pisidium in 2004, when R. philippinarum and M. incongrua had became scarce due to naticid predation. This species-selective predation affected bivalve community structure, and caused disagreements in taxonomic composition and species’ rank-order abundance between the living bivalve community and the assemblage of dead shells. This approach (live–dead analysis), frequently used in paleoecological research, is a conservative tool to identify impacts of an alien predator on community structure. When sample size is sufficient, frequency of predatory drill holes in preferred prey species is likely to reflect predation intensity.     

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.     

Predator size interacts with habitat structure to determine the allometric scaling of the functional response

While both predator body size and prey refuge provided by habitat structure have been established as major factors influencing the functional response (per capita consumption rate as a function of prey density), potential interactions between these factors have rarely been explored. Using a crab predator (Panopeus herbstii) – mussel prey (Brachidontes exustus) system, we examined the allometric scaling of the functional response in oyster (Crassostrea virginica) reef habitat, where crevices within oyster clusters provide mussels refuge from predation. A field survey of mussel distribution showed that mussels attach closer to the cluster periphery at high mussel density, indicating the potential for saturation of the refuge. In functional response experiments, the consumption rate of large crabs was depressed at low prey density relative to small crabs, while at high prey density the reverse was true. Specifically, the attack rate coefficient and handling time both decreased non‐linearly with crab size. An additional manipulation revealed that at low prey densities, the ability of large crabs to maneuver their claws and bodies to extract mussels from crevices was inhibited relative to small crabs by the structured habitat, reducing their attack rate. At high prey densities, crevices were saturated, forcing mussels to the edge of clusters where crabs were only limited by handling time. Our study illuminates a potentially general mechanism where the quality of the prey refuge provided by habitat structure is dependent on the relative size of the predator. Thus anthropogenic influences that alter the natural crab size distribution or degrade reef habitat structure could threaten the long‐term stability of the crab –mussel interaction in reefs.     

Drilling predation on the clypeasteroid echinoid Echinocyamus pusillus from the Mediterranean Sea (Giglio,Italy)

Drilling predation of cassid gastropods (tonnacean) on echinoids is common in marine environments but is rarely documented. Tests of the minute clypeasteroid Echinocyamus pusillus OF Müller (1776) were collected from the Mediterranean Sea (Isola del Giglio, Italy). Besides general morphology, features pertaining to the morphology and distribution of predatory boreholes were examined. Furthermore, borehole frequencies among different samples sites were compared. Cassid gastropods are assumed to be the originators of the boreholes. A total of 1061 tests were analysed for drilling rates with 15.3% drilled with predominantly single boreholes. The borehole morphology is strongly affected by the microstructure of the skeleton; the borehole outline is irregular if drilled within areas where ambulacral pores are present. Vertical borehole morphology is influenced by stereom density. The size frequencies of non-drilled and drilled specimens show significant differences. Comparisons of borehole size with test size show only a low correlation between predator and prey size. The distribution of boreholes shows a high site selectivity of the predator for the aboral side of the test and the petalodium.     

Phenotypic plasticity in oysters (Crassostrea virginica) mediated by chemical signals from predators and injured prey

Prey organisms reduce predation risk by altering their behavior, morphology, or life history. Avoiding or deterring predators often incurs costs, such as reductions in growth or fecundity. Prey minimize costs by limiting predator avoidance or deterrence to situations that pose significant risk of injury or death, requiring them to gather information regarding the relative threat potential predators pose. Chemical cues are often used for risk evaluation, and we investigated morphological responses of oysters (Crassostrea virginica) to chemical cues from injured conspecifics, from heterospecifics, and from predatory blue crabs (Callinectes sapidus) reared on different diets. Previous studies found newly settled oysters reacted to crab predators by growing heavier, stronger shells, but that adult oysters did not. We exposed oysters at two size classes (newly settled oyster spat and juveniles ~2.0 cm) to predation risk cue treatments including predator or injured prey exudates and to seawater controls. Since both of the size classes tested can be eaten by blue crabs, we hypothesized that both would react to crab exudates by producing heavier, stronger shells. Oyster spat grew heavier shells that required significantly more force to break, an effective measure against predatory crabs, when exposed to chemical exudates from blue crabs as compared to controls. When exposed to chemical cues from injured conspecifics or from injured clams (Mercenaria mercenaria), a sympatric bivalve, shell mass and force were intermediate between predator treatments and controls, indicating that oysters react to injured prey cues but not as strongly as to cues released by predators. Juvenile oysters of ~ 2.0 cm did not significantly alter their shell morphology in any of the treatments. Thus, newly settled oysters can differentiate between predatory threats and adjust their responses accordingly, with the strongest responses being to exudates released by predators, but oysters of 2.0 cm and larger do not react morphologically to predatory threats.     

A note on exceptionally high confamilial naticid drilling frequency on Natica gualteriana from the Indian subcontinent

Although common, confamilial naticid predation intensity was not very high in the geological record. Here, we gathered modern confamilial predation data from the Indian coasts and showed that confamilial naticid predation on a naticid species, Natica gualteriana, is exceptionally high at Chandipur, one of our studied areas. We studied the different aspects of confamilial predation from the Indian coasts and showed that the predators in Chandipur were highly efficient as evident from high drilling frequency (DF), site stereotypy and low prey effectiveness. Unusually high DF on N. gualteriana may be attributed to its new arrival in Chandipur where it faced competitive elimination through predation by sympatric naticid predators. Reports of failed invasion are rare. Natica gualteriana is a small invader and therefore its invasion success is threatened by resident populations of large species that extensively drill on young individuals of N. gualteriana just to break the bottleneck of their own offspring from competition.     

Evolutionary trends within bivalve prey of chesapeake group naticid gastropods

Five genera of Miocene bivalves evolved antipredatory adaptations in response to predation by drilling naticid gastropods. I examined the evolution of two traits affecting predator‐prey interaction, prey shell thickness (TH) and internal volume (IV). Thickness controls predation costs by determining drilling time, and internal volume influences the benefit derived by the predator.

Internal volume showed no consistent pattern of temporal change among the taxa studied. IV fluctuated nondirectionally during the history of most genera, though both increasing and decreasing trends occurred within species ofAstarte. In contrast, all five genera exhibited significant thickness increases (from 8–157%) during the three‐million‐year interval. Both gradual intraspecific and interspecific directional changes occurred. Taxa with the greatest predation intensities displayed the most change, suggesting that predation selected for the thickness increases. Increased thickness apparently reduced predation; a significant negative correlation between TH and predation intensity occurred within four of the bivalve genera. Improvement of predator capabilities apparently did not keep pace with increased antipredatory morphologic adaptations over the interval studied.     

Avoidance of drilled gastropod shells by the hermit crab Pagurus longicarpus at Nahant, Massachusetts

Most hermit crabs depend on empty gastropod shells for shelter; competition for appropriate shells is often severe. This study determined whether shells that have been drilled by naticid gastropods are suitable for occupancy by the hermit crab Pagurus longicarpus. Differences in the characteristics of empty shells and those occupied by hermit crabs were assessed at two adjacent field sites in Nahant, Massachusetts. Drilling damage was far more frequent in empty gastropod shells than in shells occupied by hermit crabs, suggesting that individuals of P. longicarpus avoid drilled shells. They did not appear to avoid shells with other forms of damage. Laboratory experiments confirmed that these hermit crabs preferentially chose intact shells over drilled shells, even when the intact shells offered were most suitable for crabs half the weight of those tested. Final shell choices were generally made within 1 h. The hermit crabs apparently discriminated between intact and drilled shells based on tactile cues, since crabs kept in the dark showed the same preference for intact shells. The hermit crabs strongly avoided, to nearly the same extent, artificially drilled shells, naturally drilled shells, and shells with holes artificially drilled on the opposite side of the shell from where they would normally be located. Possible selective forces causing P. longicarpus to show such strong behavioral avoidance of drilled shells include increased vulnerability of crabs in drilled shells to osmotic stress, predation, and eviction by conspecifics.     

Inducible defenses in Olympia oysters in response to an invasive predator

The prey naiveté hypothesis suggests that native prey may be vulnerable to introduced predators because they have not evolved appropriate defenses. However, recent evidence suggests that native prey sometimes exhibit induced defenses to introduced predators, as a result of rapid evolution or other processes. We examined whether Olympia oysters (Ostrea lurida) display inducible defenses in the presence of an invasive predator, the Atlantic oyster drill (Urosalpinx cinerea), and whether these responses vary among oyster populations from estuaries with and without this predator. We spawned oysters from six populations distributed among three estuaries in northern California, USA, and raised their offspring through two generations under common conditions to minimize effects of environmental history. We exposed second-generation oysters to cue treatments: drills eating oysters, drills eating barnacles, or control seawater. Oysters from all populations grew smaller shells when exposed to drill cues, and grew thicker and harder shells when those drills were eating oysters. Oysters exposed to drills eating other oysters were subsequently preyed upon at a slower rate. Although all oyster populations exhibited inducible defenses, oysters from the estuary with the greatest exposure to drills grew the smallest shells suggesting that oyster populations have evolved adaptive differences in the strength of their responses to predators. Our findings add to a growing body of literature that suggests that marine prey may be less likely to exhibit naiveté in the face of invasive predators than prey in communities that are more isolated from native predators, such as many freshwater and terrestrial island ecosystems.     

Reappraising the early evidence of durophagy and drilling predation in the fossil record: implications for escalation and the Cambrian Explosion

The Cambrian Explosion is arguably the most extreme example of a biological radiation preserved in the fossil record, and studies of Cambrian Lagerstätten have facilitated the exploration of many facets of this key evolutionary event. As predation was a major ecological driver behind the Explosion – particularly the radiation of biomineralising metazoans – the evidence for shell crushing (durophagy), drilling and puncturing predation in the Cambrian (and possibly the Ediacaran) is considered. Examples of durophagous predation on biomineralised taxa other than trilobites are apparently rare, reflecting predator preference, taphonomic and sampling biases, or simply lack of documentation. The oldest known example of durophagy is shell damage on the problematic taxon Mobergella holsti from the early Cambrian (possibly Terreneuvian) of Sweden. Using functional morphology to identify (or perhaps misidentify) durophagous predators is discussed, with emphasis on the toolkit used by Cambrian arthropods, specifically the radiodontan oral cone and the frontal and gnathobasic appendages of various taxa. Records of drill holes and possible puncture holes in Cambrian shells are mostly on brachiopods, but the lack of prey diversity may represent either a true biological signal or a result of various biases. The oldest drilled Cambrian shells occur in a variety of Terreneuvian‐aged taxa, but specimens of the ubiquitous Ediacaran shelly fossil Cloudina also show putative drilling traces. Knowledge on Cambrian shell drillers is sorely lacking and there is little evidence or consensus concerning the taxonomic groups that made the holes, which often leads to the suggestion of an unknown ‘soft bodied driller’. Useful methodologies for deciphering the identities and capabilities of shell drillers are outlined. Evidence for puncture holes in Cambrian shelly taxa is rare. Such holes are more jagged than drill holes and possibly made by a Cambrian ‘puncher’. The Cambrian arthropod Yohoia may have used its frontal appendages in a jack‐knifing manner, similar to Recent stomatopod crustaceans, to strike and puncture shells rapidly. Finally, Cambrian durophagous and shell‐drilling predation is considered in the context of escalation – an evolutionary process that, amongst other scenarios, involves predators (and other ‘enemies’) as the predominant agents of natural selection. The rapid increase in diversity and abundance of biomineralised shells during the early Cambrian is often attributed to escalation: enemies placed selective pressure on prey, forcing phenotypic responses in prey and, by extension, in predator groups over time. Unfortunately, few case studies illustrate long‐term patterns in shelly fossil morphologies that may reflect the influence of predation throughout the Cambrian. More studies on phenotypic change in hard‐shelled lineages are needed to convincingly illustrate escalation and the responses of prey during the Cambrian.