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.     

Pathologies of non‐marine bivalve shells from the Late Triassic of Poland

Gorzelak, P., Nied?wiedzki, G. & Skawina, A. 2010: Pathologies of non‐marine bivalve shells from the Late Triassic of Poland. Lethaia, Vol. 43, pp. 285–289. Shells of Late Triassic non‐marine bivalves from Lisowice (Lipie ?l?skie clay pit, southern Poland), which co‐occur with remains of several vertebrate taxa (mammal‐like reptiles, carnivorous dinosaurs, pterosaurs, temnospondyl amphibians, hybodont sharks, dipnoan and ganoid fish), bear evidence of pathologies. Distribution, dimension and shape of some of these injuries (radiate tooth marks) closely match the dental morphology of lungfish (here probably represented by the genus Ceratodus). Thus, we interpret these pathologies as evidence of unsuccessful predatory attack on bivalves by this fish. This interpretation is also consistent with modern examples of such behaviour among lungfish. The feasibility that other culprits caused other pathologies (shell scarring and wedges) on the bivalves analysed is also discussed. Discovery of these traces constitutes important evidence of predator–prey interaction, which provides ‘fingerprints’ of trophic structure within this Late Triassic freshwater ecosystem. □Freshwater bivalves, lungfish, pathologies, predation, Triassic.     

The earliest post-paleozoic freshwater bivalves preserved in coprolites from the karoo basin, South Africa

Background

Several clades of bivalve molluscs have invaded freshwaters at various times throughout Phanerozoic history. The most successful freshwater clade in the modern world is the Unionoida. Unionoids arose in the Triassic Period, sometime after the major extinction event at the End-Permian boundary and are now widely distributed across all continents except Antarctica. Until now, no freshwater bivalves of any kind were known to exist in the Early Triassic.

Principal Findings

Here we report on a faunule of two small freshwater bivalve species preserved in vertebrate coprolites from the Olenekian (Lower Triassic) of the Burgersdorp Formation of the Karoo Basin, South Africa. Positive identification of these bivalves is not possible due to the limited material. Nevertheless they do show similarities with Unionoida although they fall below the size range of extant unionoids. Phylogenetic analysis is not possible with such limited material and consequently the assignment remains somewhat speculative.

Conclusions

Bivalve molluscs re-invaded freshwaters soon after the End-Permian extinction event, during the earliest part of the recovery phase during the Olenekian Stage of the Early Triassic. If the specimens do represent unionoids then these Early Triassic examples may be an example of the Lilliput effect. Since the oldest incontrovertible freshwater unionoids are also from sub-Saharan Africa, it is possible that this subcontinent hosted the initial freshwater radiation of the Unionoida. This find also demonstrates the importance of coprolites as microenvironments of exceptional preservation that contain fossils of organisms that would otherwise have left no trace.     

Shell decay rates of native and alien freshwater bivalves and implications for habitat engineering

1. Spent shells of bivalves can provide habitat for other organisms, as well as playing important roles in biogeochemical cycles. The amount of spent shell material that will accumulate at a site depends on rates of both shell production and decay, although the latter is rarely considered. 2. We measured the instantaneous decay rates of four species of freshwater bivalves across a range of sites in south‐eastern New York, and found that rates varied by more than 500‐fold across sites and species. 3. Differences in decay rates were related to water chemistry (Ca, pH, dissolved inorganic C), the presence of a current, and the size of the bivalve shell. 4. Combining these decay rates with estimates of shell production derived from the literature, we conclude that the Unionidae, Corbicula, and Dreissena are all capable of producing large accumulations (>10 kg dry mass m⁻²) of spent shells, while members of the Sphaeriidae probably rarely will produce such large accumulations. 5. Hence the replacement of native unionid bivalves by the alien Corbicula and Dreissena may have little effect on standing stocks of spent shells, unless the aliens invade sites where unionids are scarce or absent.     

New Trigonioidoidea (Bivalvia; Unionoida) from the Early Cretaceous of Spain

Abstract: Two taxa belonging to the Trigonioidoidea (Order Unionoida) are described from the Early Cretaceous of Spain. Nippononaia (Paranippononaia) camerana subgen. et sp. nov. is described from the Aptian of the Cameros Basin of the north‐west Iberian Range. Subnippononaia fordi Barker et al., 1997 is described from new material from the Calizas de la Huérguina Formation (Late Barremian) from Las Hoyas and Buenache de la Sierra, Cuenca Province, and Subnippononaia is raised to generic status. The stratigraphy and palaeoenvironment of the Cornago G exposure of the Enciso Group where Nippononaia (Paranippononaia) camerana originates is interpreted as a fresh to brackish water system.     

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.     

Identifying mammalian predators from bite marks: a tool for focusing wildlife protection

Dead Sooty Shearwater, Puffinus griseus, chicks and adults were collected from seven colonies on South Island, New Zealand in the 1993–96 breeding seasons. An estimated 97% of 118 deaths were from predation. Thirty‐four definite predator bite pairs were identified on 27 carcasses. Twenty‐one (78%) of the carcasses had bite pairs with intercanine distances < 9.5 mm which suggests that Stoats (Mustela erminea) were the principal predators. One chick was killed by a feral House Cat (Felis catus), and it is likely that feral Ferrets (M. furo) were responsible for a proportion of the deaths. Nearly three quarters of definite Stoat bite pairs were identified in the head region. The analyses of bite marks offers cheap and statistically reliable identification of predators provided carcasses are collected fresh and flesh is removed to examine tooth punctures in bone.     

Predation on exotic zebra mussels by native fishes: effects on predator and prey

SUMMARY 1. Exotic zebra mussels, Dreissena polymorpha, occur in southern U.S. waterways in high densities, but little is known about the interaction between native fish predators and zebra mussels. Previous studies have suggested that exotic zebra mussels are low profitability prey items and native vertebrate predators are unlikely to reduce zebra mussel densities. We tested these hypotheses by observing prey use of fishes, determining energy content of primary prey species of fishes, and conducting predator exclusion experiments in Lake Dardanelle, Arkansas. 2. Zebra mussels were the primary prey eaten by 52.9% of blue catfish, Ictalurus furcatus; 48.2% of freshwater drum, Aplodinotus grunniens; and 100% of adult redear sunfish, Lepomis microlophus. Blue catfish showed distinct seasonal prey shifts, feeding on zebra mussels in summer and shad, Dorosoma spp., during winter. Energy content (joules g⁻¹) of blue catfish prey (threadfin shad, Dorosoma petenense; gizzard shad, D. cepedianum; zebra mussels; and asiatic clams, Corbicula fluminea) showed a significant species by season interaction, but shad were always significantly greater in energy content than bivalves examined as either ash-free dry mass or whole organism dry mass. Fish predators significantly reduced densities of large zebra mussels (>5 mm length) colonising clay tiles in the summers of 1997 and 1998, but predation effects on small zebra mussels (≤5 mm length) were less clear. 3. Freshwater drum and redear sunfish process bivalve prey by crushing shells and obtain low amounts of higher-energy food (only the flesh), whereas blue catfish lack a shell-crushing apparatus and ingest large amounts of low-energy food per unit time (bivalves with their shells). Blue catfish appeared to select the abundant zebra mussel over the more energetically rich shad during summer, then shifted to shad during winter when shad experienced temperature-dependent stress and mortality. Native fish predators can suppress adult zebra mussel colonisation, but are ultimately unlikely to limit population density because of zebra mussel reproductive potential.     

Quantifying phenotypic gradients in freshwater snails: a case study in Lithasia (Gastropoda: Pleuroceridae)

Many authors have described a pattern of morphological variation in freshwater bivalves where shells taken from lentic and lotic environments, or headwaters and main stem reaches, appear to exhibit phenotypic gradients in size and shape. For example, mussels taken from headwater reaches tend to possess smooth, less inflated shells compared to the more obese, sculptured individuals downstream. Others observed similar relationships in certain freshwater gastropods, but this variation has not been quantified nor its existence explained in an ecological or evolutionary context. Geometric morphometrics indicated freshwater snails shells from the pleurocerid genus Lithasia from the Duck River, Tennessee, USA, show phenotypic gradients similar to those in freshwater mussels. Shells from upstream areas were narrow and less sculptured on the posterior portions of their body whorls, while downstream shells were more inflated and possessed significantly more sculpture. This phenotypic variation may reduce predation or damage due to dislodging. The nature of the observed plasticity suggests an unidirectional environment similar to that proposed by the river continuum concept. Handling editor: K. Martens     

Morphological and ecological shifts in a land snail caused by the impact of an introduced predator

Introduced predators have become major threats to native animal species in oceanic islands. A number of studies have shown that alien predators have caused serious extinctions of island endemics. However, little attention has been paid to the evolutionary impacts of alien predators on native species. The present study shows that predation by black rats, Rattus rattus, has resulted in ecological and morphological changes in the land snail Mandarina anijimana from the island of Anijima in the Ogasawara archipelago. The frequency of empty predated shells has increased over the past 17–19 years in southern areas of the island. The shells of these snails were found to be significantly higher, smaller and darker in the survey in 2006 than in the survey in 1987–1989 performed in central and southern parts of Anijima, where predation by Rattus was serious. M. anijimana were formerly restricted to shallow broad-leaved litter, whereas they are currently found in deep palm litter, where predation pressure from Rattus may be lower. This suggests that increased predation pressure by Rattus has changed the habitat use of M. anijimana. The close association between shell morphology and habitat use of Mandarina species suggests that the habitat shift induced by the predation of Rattus has caused these changes in the shell morphology of M. anijimana over a period of 17–19 years.     

Mid-Neolithic Exploitation of Mollusks in the Guanzhong Basin of Northwestern China: Preliminary Results

Mollusk remains are abundant in archaeological sites in the Guanzhong Basin of Northwestern China, providing good opportunities for investigations into the use of mollusks by prehistoric humans. Here we report on freshwater gastropod and bivalve mollusks covering the time interval from about 5600 to 4500 cal. yrs BP from sites of Mid-Late Neolithic age. They are identified as Cipangopaludina chinensis and Unio douglasiae, both of which are currently food for humans. The shells are well preserved and have no signs of abrasion. They are all freshwater gastropods and bivalves found in pits without water-reworked deposits and have modern representatives which can be observed in rivers, reservoirs, and paddy fields in the studied region. Mollusk shells were frequently recovered in association with mammal bones, lithic artifacts, and pottery. These lines of evidence indicate that the mollusks are the remains of prehistoric meals. The mollusk shells were likely discarded into the pits by prehistoric humans after the flesh was eaten. However, these mollusk remains may not have been staple food since they are not found in large quantities. Mollusk shell tools and ornaments are also observed. Shell tools include shell knives, shell reaphooks and arrowheads, whereas shell ornaments are composed of pendants and loops. All the shell tools and ornaments are made of bivalve mollusks and do not occur in large numbers. The finding of these freshwater mollusk remains supports the view that the middle Holocene climate in the Guanzhong Basin may have been warm and moist, which was probably favorable to freshwater mollusks growing and developing in the region.     

The isotopic biosignatures of photo‐ vs. thiotrophic bivalves: are they preserved in fossil shells?

Symbiont‐bearing and non‐symbiotic marine bivalves were used as model organisms to establish biosignatures for the detection of distinctive symbioses in ancient bivalves. For this purpose, the isotopic composition of lipids (δ¹³C) and bulk organic shell matrix (δ¹³C, δ³⁴S, δ¹⁵N) from shells of several thiotrophic, phototrophic, or non‐symbiotic bivalves were compared (phototrophic: Fragum fragum, Fragum unedo, Tridacna maxima; thiotrophic: Codakia tigerina, Fimbria fimbriata, Anodontia sp.; non‐symbiotic: Tapes dorsatus, Vasticardium vertebratum, Scutarcopagia sp.). ?¹³C values of bulk organic shell matrices, most likely representing mainly original shell protein/chitin biomass, were depleted in thio‐ and phototrophic bivalves compared to non‐symbiotic bivalves. As the bulk organic shell matrix also showed a major depletion of δ¹⁵N (down to –2.2 ‰) for thiotrophic bivalves, combined δ¹³C and δ¹⁵N values are useful to differentiate between thio‐, phototrophic, and non‐symbiotic lifestyles. However, the use of these isotopic signatures for the study of ancient bivalves is limited by the preservation of the bulk organic shell matrix in fossils. Substantial alteration was clearly shown by detailed microscopic analyses of fossil (late Pleistocene) T. maxima and Trachycardium lacunosum shell, demonstrating a severe loss of quantity and quality of bulk organic shell matrix with time. Likewise, the composition and δ¹³C‐values of lipids from empty shells indicated that a large part of these compounds derived from prokaryotic decomposers. The use of lipids from ancient shells for the reconstruction of the bivalve's life style therefore appears to be restricted.     

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.     

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.     

Ventral bite marks in Mesozoic ammonoids

Reports on the predators of ammonoids are rare, although ammonoids were abundant and diverse invertebrates in many Paleozoic and Mesozoic marine ecosystems. Most previous work on lethal ammonoid predation has focused on (sub)circular tooth marks which resulted from fish and mosasaur attacks. In the present study we discuss a relatively common type of bite mark in ammonoid shells, the ‘ventral bite mark’. This typically occurs in a restricted position on the ventral side of the outer body chamber whorl and does not affect either the aperture or the phragmocone. Ammonoid specimens revealing ventral bite marks used in this study were collected from a wide range of strata which range in age from the Lower Jurassic to the uppermost Cretaceous (close to the Cretaceous–Paleogene boundary). These ventral bite marks are absent in the Paleozoic collections studied. The vast majority of ventral bite marks are situated at the end of the body chamber, close to the phragmocone. This is interpreted as the result of predatory attacks on the back or blind side of ammonoids in their living position. The predators aimed for the vital parts and muscle attachments to obtain the edible soft tissues. The agents for most of the ventral bite marks to ammonoids are probably coleoid cephalopods (especially teuthoids) and predatory fishes to a lesser extent.     

What makes a good home for hermits? Assessing gastropod shell density and relative strength

The survival and reproductive success of hermit crabs is intrinsically linked to the quality of their domicile shells. Because damaged or eroded shells can result in greater predation, evaluating shell structure may aid our understanding of population dynamics. We assessed the structural attributes of Cerithium atratum shells through assessments of (a) density using a novel approach involving computed tomography and (b) tolerance to compressive force. Our goal was to investigate factors that may influence decision making in hermit crabs, specifically those that balance the degree of protection afforded by a shell (i.e. density and strength) with the energetic costs of carrying such resources. We compared the density and relative strength (i.e. using compression tests) of shells inhabited by live gastropods, hermit crabs (Pagurus criniticornis) and those found empty in the environment. Results failed to show any relationship between density and shell size, but there was a notable effect of shell density among treatment groups (gastropod/empty/hermit crab). There was also a predictable effect of shell size on maximum compressive force, which was consistent among occupants. Our results suggest that hermit crabs integrate multiple sources of information, selecting homes that while less dense (i.e. reducing the energy costs of carrying these resources), still offer sufficient resistance to compressive forces (e.g. such as those inflicted by shell-breaking predators). Lastly, we show that shell size generally reflects shell strength, thus explaining the motivation of hermit crabs to search for and indeed fight over the larger homes.     

Differential vulnerability to predation in two sympatric whelks is mediated by juvenile traits

In many taxa, initial differences in offspring size play an important role in mediating subsequent performance; however, the consequences of interspecific variation in size for the performance of co‐occurring taxa have been rarely examined. We used the whelks Cominella virgata and C. maculosa, which co‐occur on rocky shores throughout their life cycles, to examine the vulnerability of early life‐stages to native predators under controlled laboratory conditions. Among all the predators evaluated (the cushion sea star Patiriella spp., the olive rockfish Acanthoclinus fuscus, the oyster borer snail Haustrum scobina, the smooth shore crab Cyclograpsus lavauxi, and the pebble crab Heterozius rotundifrons), hatchlings of both species (C. virgata: ~3 mm shell length [SL] and C. maculosa: ~1.5 mm SL) were especially vulnerable to the smooth shore crab Cy. lavauxi, the only potential predator in which mortality was greater than in the control treatment. Small shore crabs (~8 mm carapace width [CW]) were unable to eat hatchlings of either whelk species, whereas medium and large shore crabs (~12 and ~18 mm CW, respectively) consumed hatchlings of both prey species. Hatchlings of C. virgata were less vulnerable to predation by medium crabs than large ones, and those of C. maculosa were equally vulnerable to both sizes of crabs. In hatchlings of both prey species, shell length and shell thickness increased over time. Two months after hatching, only individuals of C. virgata had reached a size refuge from predation. Our results show that interspecific vulnerability to predators can be mitigated by larger sizes and thicker shells at hatching; nonetheless, our results also suggest that other species‐specific factors, such as juvenile growth rate, may also play key roles in determining the vulnerability of hatchling and juvenile snails to shell‐crushing predators.     

Anatomy of a cline: dissecting anti‐predatory adaptations in a marine gastropod along the U.S. Atlantic coast

The scope of anti‐predatory adaptation is expected to be greater in warm than in cold environments. High temperatures lower the costs associated with the production and maintenance of energetically expensive traits and enable ecological interactions to intensify. We tested this hypothesis by characterizing the expression of anti‐predatory morphology within a marine gastropod species (the knobbed whelk Busycon carica) over a large (> 1400 km) geographic area that spans more than 10°C annual temperature variation. We also conducted experimental predation studies with a powerful durophagous predator, the stone crab Menippe, to verify the anti‐predatory advantages of a heavily ornamented shell morphology (e.g. increased thickness, pronounced spines), and we used repair scar data to assess clinal variation in selective pressure from predators. We predicted that repair scar rates would be greatest in warm southernmost latitudes, and that expression of energetically costly anti‐predatory morphology would peak in concert with elevated predation pressures. Experiments confirmed that whelks with energetically costly, heavily ornamented shells had higher survivorship rates than those with weakly ornamented shells. As predicted, we also found that the expression of anti‐predatory traits was greatest in the southern part of B. carica's range. After standardizing shells for size, shape, and exposure time to enemies, repair scar rates also peaked to the south. Taken together, these results suggest that the expression of anti‐predatory traits along the geographic cline is governed by the interaction of two selective factors: temperature and predation, with the former acting as the ultimate control on the scope of adaptation both by escalating predation pressure in the southern part of B. carica's range and by physically limiting (to the north) and facilitating (to the south) the production of anti‐predatory traits. Feedbacks between temperature and predation thus causally interact to enable and drive, respectively, the observed geographic cline in energy‐intensive anti‐predatory shell traits.     

Opisoma excavatum Boehm,a Lower Jurassic photosymbiotic alatoform‐chambered bivalve

Alatoform bivalves are a polyphyletic group characterized by antero‐posteriorly compressed shells and a ventro‐lateral wing originating from a tight fold of the shell wall. This bizarre shell morphology is interpreted as an adaptation for algal photosymbiosis in heliophilous bivalves. The group contains the living heart cockle Corculum together with four extinct genera ranging in age from the Permian to the Jurassic. The Jurassic alatoform bivalve is Opisoma, which has an aragonitic shell that is divided into two regions, both with different functions: one for stabilization, the other for hosting symbionts. The dorsal part of the shell is massive and played the stabilization role. The ventral part has a very thin and fragile shell that permitted the transmission of light into the internal tissues harbouring photosymbionts. The morphology of this delicate ventral part has thus far remained obscure, due to lack of preservation. Accumulations of Opisoma excavatum Boehm with exquisitely preserved shells containing the fragile winged ventral parts are common within the Pliensbachian shallow‐water, lagoonal carbonate succession of the Rotzo Formation of northern Italy. The wings have internal curved chambers limited by septa parallel to the wing edge. The shell of the ventral part consists of irregular fibrous prismatic and homogeneous structures which progressively infill the chambers. As the chambered wings are analogous structures among alatoform bivalves, they are no longer considered a taxonomic character. According to the observed shell orientation in the field and the consequent organization of the soft parts, Opisoma had an opisthogyrate shell.     

Signs of predation in the Middle Jurassic of south-central Poland: evidence from echinoderm taphonomy

Distinct faunal aggregates are described from the Middle Jurassic (uppermost Bajocian/lowermost Bathonian and Middle Bathonian) clay deposits of Częstochowa area, south-central Poland. These aggregates are composed of molluscs (scaphopods, gastropods, bivalves, ammonites and belemnites), articulate brachiopods and echinoderms (asteroids, crinoids and echinoids). A large percentage of the fossils, especially bivalves, are fragmented, but some fossils are complete. Although most of the fossils are crushed and fragmented, they are still identifiable to at least the genus level. Thorough statistical analysis of taphonomic features indicates that the preservation of asteroid marginal plates is distinct from the ossicles derived from the host clays. The high frequency of bite marks and the good state of preservation suggest that the accumulations are the products of predation activities and most probably are the effect of regurgitation. Taking into account the rich and diverse fauna, the predator was a bottom-feeding generalist. Possible predators include palaeospinacid sharks, a tooth of which was collected from the same bedding surface, but not associated with regurgitated remains. Although the bite marks on the asteroid ossicles point to sharks as potential producers of regurgitates, other vertebrates, like durophagous pycnodontiform fish, cannot be excluded.