Host‐specific acrothoracid barnacles on Middle Devonian platyceratid gastropods

Borings, attributed to acrothoracic barnacles, occur on the platyceratid gastropod Naticonema lineatum (Conrad) from the Middle Devonian Hamilton Group of western New York and rarely in specimens as old as the Early Devonian. These latter are the oldest known acrothoracid borings are in the fossil record. The borings are consistently developed as laterally compressed, inequilateral pouches exclusively on these gastropods, commonly as dense infestations.

Naticonema shells yielding borings typically occur associated with partially articulated qrinoid remains, and they are sometimes found attached to crinoids in a manner similar to coprophagous Platyceras. In addition to barnacle borings, Naticonema shells often bear thin encrustations of bryozoans which are usually perforated by these borings but sometimes also overgrow them. Barnacles bored live hosts; gastropods prevented shell penetration by producing cyst‐like secondary secretions of calcite beneath acrothoracid boreholes.

The relative antiquity of these borings and their association with coprophagous platyceratids makes their discovery particularly significant in revealing aspects of the early ecology of barnacles. Attachment to the host commensal gastropods was one of the first successful life modes of these crustaceans prior to their later diversification to other habitats. Mississippian and Pennsylvanian occurrences of similarly bored gastropods demonstrate continuity of the barnacle‐gastropod‐crinoid ecological association from the Middle to Late Paleozoic.     

<Emphasis Type="Italic">Toarctocera</Emphasis> (Gastropoda,Aporrhaidae): a new genus from the Jurassic (Toarcian/Aalenian) of South Germany and the early evolutionary history of the family Aporrhaidae

Toarctocera, a new gastropod genus of the family Aporrhaidae (Stromboidea) is described for Rostellaria subpunctata. It occurs in Central Europe (S Germany, N Switzerland, and France) and is commonly one of the most abundant gastropods in Late Toarcian/Early Aalenian soft bottom faunas. Toarctocera subpunctata has extremely elongated apertural spines; this has been interpreted as an adaptation to extremely soft-bottom conditions (snowshoe strategy). The new genus represents one of the earliest certain aporrhaids (Stromboidea). Aporrhaids are probably as old as Late Triassic, and possible Triassic and Early Jurassic species are discussed. Spiniloma is proposed as a nomen novum in order to replace the homonym Spinigera and consequently the family Spinilomatinae replaces the Spinigeridae. The sister group of Stromboidea still needs to be identified. A previously suggested sister group relationship with the Heteropoda is seen as unlikely.     

Patellogastropoda and Vetigastropoda (Mollusca,Gastropoda) from the marine Jurassic of Patagonia,Argentina

Gastropod faunas from the Early Jurassic (Late Pliensbachian–Early Toarcian) marine deposits of Chubut Province, Argentina, are described from Lomas Occidentales, Cerro La Trampa and Puesto Currumil localities, representing eight species, three of them new. These are Scurriopsis? sp., Chartronella gradata sp. nov., Calliotropis? sp., Pleurotomaria sp., Leptomaria sp., Hamusina? wahnishae sp. nov., Colpomphalus musacchioi sp. nov. and Jurassiphorus? cf. triadicus Haas. The gastropod assemblage reported here testifies paleobiogeographical connections with other coeval gastropod associations from the western Tethys. However, Chartronella, Hamusina and Jurassiphorus may represent survivors of Triassic associations, considering the ancient seaway from Peru as the most plausible hypothesis for biotic exchange of these faunas during the Late Triassic–Early Jurassic boundary. An abundant and diverse invertebrate fauna such as corals, echinoderms, cephalopods, brachiopods, bivalves and other gastropods found in association with the gastropods described here characterises a shallow marine environment for the gastropod-bearing rocks.http://www.zoobank.org/urn:lsid:zoobank.org:pub:7B8EAFC0-3AC0-4F91-97A5-22AAC6A19909     

Pliensbachian gastropods from Venetian Southern Alps (Italy) and their palaeobiogeographical significance

Abstract: The Pliensbachian gastropods described by De Toni in 1912, coming from an isolated boulder at the foot of Mt Vedana (eastern margin of Trento Platform, Venetian Southern Alps, Italy) are revised. The fauna consists of 13 species representing nine families and eight superfamilies. Despite the low number of species, the assemblage represents the most diverse Early Jurassic gastropod fauna known for the Venetian Southern Alps. The boulder yielding the material was thought to derive from the upper part of the Early Jurassic Calcari Grigi Group, a carbonate platform unit extensively cropping out in the Mt Vedana area. The sedimentological analysis indicates a prevalently bioclastic wackstone‐floatstone, reflecting a lime‐muddy deposit undergone to an early consolidation. This and the high content of ammonoids, which is unusual for the Calcari Grigi Group, are typical aspects of a condensed pelagic sediment, presumably a fissure filling at the top of the carbonate platform succession. Palaeobiogeographical comparisons show that the fauna is composed of species occurring exclusively in pelagic limestones of the western Tethys. By contrast, it shows no relationships with the coeval faunas of the adjacent Trento platform and of the other western Tethyan carbonate platforms. These lines of evidence and the facies analysis would testify to the Pliensbachian drowning of the eastern margin of the Trento platform. In the wider context of the palaeobiogeographical history of Early Jurassic western Tethyan gastropods, the species from Vedana belong to a faunal stock which is typical for pelagic, mainly postdrowning sediments. Thus, appearance and diffusion in space and time of this stock were probably regulated by the direction, rate and pattern of the Neotethyan rifting. A new subgenus, Proarcirsa (Schafbergia) subgen. nov., and three new species, namely Ataphrus (Ataphrus) cordevolensis sp. nov., Guidonia pseudorotula sp. nov. and Proarcirsa (Schafbergia) zirettoensis sp. nov. are erected.     

Early jurassic (Upper Pliensbachian) gastropods from the Herforder Liasmulde (Bielefeld, Northwest Germany)

An Early Jurassic (Upper Pliensbachian) gastropod fauna from the Herforder Liasmulde (Herford Lias Syncline) near Bielefeld, Northwest Germany is described. This is one of the first detailed and illustrated reports of Pliensbachian gastropods from this area. Sixteen species are reported. Surface collecting yielded relatively large vetigastropods (pleurotomarioids amd trochomorphs). Bulk samples yielded abundant small caenogastropods (especiallyLevipleura blainvillei andKalchreuthia frankei) and cylindrobullinid opisthobranchs (e.g.Cylindrobullina dornend). These small species are also abundant in the Pliensbachian of South Germany (N Bavaria, Franconia). Therefore, this characteristic Pliensbachian microgastropod assemblage was widespread in Germany and probably also in other comparable marine deposits of Central Europe (e.g. the Paris Basin).      

Evolution of gastropod mitochondrial genome arrangements

Background  

Gastropod mitochondrial genomes exhibit an unusually great variety of gene orders compared to other metazoan mitochondrial genome such as e.g those of vertebrates. Hence, gastropod mitochondrial genomes constitute a good model system to study patterns, rates, and mechanisms of mitochondrial genome rearrangement. However, this kind of evolutionary comparative analysis requires a robust phylogenetic framework of the group under study, which has been elusive so far for gastropods in spite of the efforts carried out during the last two decades. Here, we report the complete nucleotide sequence of five mitochondrial genomes of gastropods (Pyramidella dolabrata, Ascobulla fragilis, Siphonaria pectinata, Onchidella celtica, and Myosotella myosotis), and we analyze them together with another ten complete mitochondrial genomes of gastropods currently available in molecular databases in order to reconstruct the phylogenetic relationships among the main lineages of gastropods.     

Potamopyrgus antipodarum(Mollusca:Hydrobiidae) in continental aquatic gastropod communities: impact of salinity and trematode parasitism

The structure of gastropod communities was examined from January to June 1999 in four sites of the streams of Mont Saint-Michel Bay along a gradient of salinity, and the occurrence of larval trematodes infecting snails was studied. Abundance and species richness of gastropods increased from polyhaline (95 snails, 1 species) to oligohaline waters (6672 snails, 6 species). Whatever the salinity, the most abundant species was Potamopyrgus antipodarum, an invasive non-indigenous species that represented 80% of the gastropods. Only one male was found in P. antipodarum populations suggesting a predominantly parthenogenetic mode of reproduction. Among 7218 gastropods collected, 1.2% were infected by larval trematodes: 5 species in Lymnaea peregra (4.4%), 4 species in Planorbis planorbis (12.0%), one echinostome in Physa acuta (0.2%), and a new species of Sanguinicola in P. antipodarum (0.5%). This is the first record of infected P. antipodarum in Europe. No parasites were found in polyhaline waters. The prevalence per host population varied from 0 to 100% depending on time of collection, salinity and host species. In the lowest-salinity site, abundance of gastropods and prevalence of trematodes were negatively correlated. The dominance of P. antipodarum in the gastropod communities is discussed in relation with euryhalinity, parthenogenesis and weak rate of parasitism.     

Anatomy and systematic affinity of Stanleya neritinoides (Smith, 1880), an enigmatic member of the thalassoid gastropod species flock in Lake Tanganyika, East Africa (Cerithioidea, Paludomidae)

The radiation of gastropods in Lake Tanganyika is an ideal system for testing competing hypotheses of species flock formation. Yet, much of the basic biology of these species remains unknown. In an ongoing effort to understand the evolution of Tanganyikan gastropods, we here describe Stanleya neritinoides. Alcohol‐preserved material of the soft parts is rare, consequently, the systematic position of the species, and a repeated suggested affinity to Tanganyicia rufofilosa, have been based primarily on features of the shell. However, features of the radula and operculum are unique and do not suggest an affinity to any other Tanganyikan species. Thus, S. neritinoides has remained a particularly poorly known and enigmatic member of the species flock. This investigation confirmed that several aspects of internal and external anatomy are shared between S. neritinoides, T. rufofilosa, and other Tanganyikan gastropods, but that S. neritinoides is unique in features of the radula and seminal receptacle. Moreover, S. neritinoides differs from T. rufofilosa in features of the foregut, midgut, hindgut, kidney, nervous system, reproductive system and reproductive strategy. These new data are inconsistent with an interpretation of identity of Stanleya and Tanganyicia. In addition, given the pervasive differences between the two, a sister‐group relationship between the two is unlikely. More precise systematic placement of S. neritinoides awaits the establishment of a phylogenetic framework for all Tanganyikan gastropods.     

Introduction,distribution, spread,and impacts of exotic freshwater gastropods in Texas

We examined the patterns of distribution, vectors of introduction, and potential ecological impacts of freshwater exotic species in Texas over the last 45 years. Currently, five species of exotic gastropods are established: channeled-type applesnail (Pomacea insularum), red-rim melania (Melanoides tuberculatus), quilted melania (Tarebia granifera), giant rams-horn snail (Marisa cornuarietis), and Chinese mysterysnail (Cipangopaludina chinensis). In contrast to the northern part of the US, where shipping appears to be the most important vector for the introduction of aquatic invasive species, aquarium and ornamental trade dominated among unintentional vectors of introduction of all freshwater exotics in Texas, resulting in different patterns of distribution, spread, and ecological impacts. The rate of spread of exotic gastropods in Texas varied from 39 waterbodies colonized over 18 years for P. insularum to only three waterbodies during last 45 years for C. chinensis. Four of five exotic gastropods were found in highly vulnerable aquifer-fed springs and rivers, which contain numerous endemic and endangered species. The fifth species, Pomacea insularum, is an agricultural pest. Potential negative ecological effects of exotic gastropods include impacts on wetlands and wetland restoration, competitive exclusion of native snails, and the introduction of exotic parasites, trematodes, which could infect fish and waterfowl, including federally protected species. Aquifer springs with stable temperature regimes are refuges for both cold and warm intolerant species. Handling editor: D. Dudgeon     

Brood care among basommatophorans: a unique reproductive strategy in the freshwater limpet snail Protancylus (Heterobranchia: Protancylidae), endemic to ancient lakes on Sulawesi, Indonesia

In molluscan taxa inhabiting marine environments oviparity and reproduction via planktonic larvae is predominant while incubation and viviparity is most frequently found in taxa inhabiting brackish or freshwater aquatic habitats. Brooding has evolved repeatedly and independently in several limnic taxa among Bivalvia and Gastropoda. However, among basommatophoran gastropods no such cases were yet known. We here report on a unique reproductive strategy involving brood care in the lacustrine freshwater limpet genus Protancylus, endemic to the ancient lakes on central Sulawesi (former Celebes), Indonesia, namely the Lake Poso and the Malili lake system, because this constitutes the first known case of this behaviour among the Basommatophora. Protancylus live exclusively as epizoans on those pachychilid gastropods of the viviparous genus Tylomelania, also a Sulawesi endemic species, that inhabit mostly soft substrates. We found that the two known species Protancylus pileolus from Lake Poso and P. adhaerens from the Malili lake system both retain gelatinous egg strings underneath their outer mantle, where up to 15 (mostly eight or nine) shelled juveniles are brooded. Nourishment is provided within the egg capsule only. Thus, brood care in Protancylus resembles the reproductive strategy found recently among pachychilid gastropods Jagora from the Philippines, but differs from euviviparous (i.e. matrotrophic) incubation among thiarid gastropods possessing a brood pouch with juveniles being nourished via a ‘pseudoplacenta’ in several taxa.     

Phenotypic plasticity in gastropod shell remodeling

A laboratory experiment was conducted with the marine gastropod Littorina littorea from three different sites in Long Island Sound to test for inducible responses to the non‐native predatory crab Hemigrapsus sanguineus. Individuals of L. littorea did not exhibit decreased shell growth in response to increased predation risk, which differs from the response seen in other species of Littorina. However, snails did exhibit slower growth in response to reduced food availability, as seen in other gastropods. Surprisingly, snails from two sites exhibited differences between the crab‐exposed and reduced food‐availability treatments in how they remodeled the thickness of different microstructural layers of shell material in apical parts of the shell. Although individuals of L. littorea did not show the typical shell thickening response seen in other gastropods, they did show differences among populations in remodeling their shell. This underlies the importance of examining how the shell is constructed and remodeled in studies of inducible defenses in gastropods, as well as the importance of examining animals from multiple populations.     

<Emphasis Type="Italic">Theba pisana</Emphasis> (Müller, 1774) (Gastropoda,Helicidae) and other alien land molluscs species in Argentina

The presence and distribution in Argentina of the invasive snail species Theba pisana is presented on the basis of a survey carried out in 10 beach resorts along the Atlantic coast of Buenos Aires Province. Additionally, complementary information and complete revision of material of other exotic terrestrial gastropods housed in museum collections was carried out, reporting herein seven new species for Argentina (Otala punctata, Hawaiia minuscula, Paralaoma servilis, Opeas goodalli, Vallonia pulchella, Vertigo ovata and Pupisoma dioscoricola), one for Colombia (Subulina octona), and two for Peru (S. octona and P. dioscoricola). At present, 42 introduced species of terrestrial gastropods have been recorded in nine countries of South America (Argentina, Bolivia, Brazil, Chile, Colombia, Paraguay, Peru, Uruguay and Venezuela). The most likely pathways for introduction of terrestrial gastropods, at least in Argentina, are horticultural development and urban and suburban transformation of original natural habitats.     

The Small-Subunit rRNA Gene Sequences of Venerids and the Phylogeny of Bivalvia

The complete nucleotide sequence of the 18S subunit of ribosomal DNA (rDNA) was determined for the venerid clams Callista chione (Pitarinae) and Venus verrucosa (Venerinae). Comparison of the new sequences with the published sequences of 1 annelid, 2 gastropods, 2 polyplacophorans, and 19 bivalves showed that when the annelids are used as outgroup the gastropods diverge from the bivalves, which form a cluster including the polyplacophorans. When the gastropods alone were compared with the bivalves, the latter split in two groups corresponding to the two subclasses of Heterodonta and Pteriomorpha. The former include two taxa that diverged early, Galeomma and Tridacna, while the Veneridae and Mactridae form two sister groups. In contrast to previous reports and in line with morphological data, the Ostreidae are included in the Pteriomorphia and form a monophyletic group. Received: 16 May 1998 / Accepted: 11 August 1998     

The chondrichthyan fauna from the Middle-Late Triassic of Guanling (Guizhou province,SW China)

The Recent volutids Cymbiola and Melo burrow in a forward direction. This agrees well with the spiral terrace pattern observed in Eocene Volutidae, thus strengthening the idea that they were forward burrowers as well. The presence of collabral terraces near the aperture in the Cassidae suggests an oblique burrowing direction, by convergence with the Nassariidae. This is confirmed by observations on living cassids. The sutural canals and associated structures of the Olividae and Seraphidae are functional in detecting whether the posterior region of the shell is buried. This confirms the idea that fossil Strombidae possessing similar features were burrowers.

In contrast with these “predictable”; observations, burrowing was observed in a few gastropods the shell morphology of which would seem to exclude such behaviour, such as Architectonica, Cymatium, Volema and Bolinus. Thus, a few morphologic criteria appear to be reliable and of general applicability in inferring burrowing habits in gastropods. At the same time, the broad variety of burrowing mechanisms and life habits of gastropods makes it unlikely that general criteria will ever be found to exclude burrowing habits on the basis of shell morphology.     

A Middle Miocene endemic freshwater mollusc assemblage from an intramontane Alpine lake (Aflenz Basin,Eastern Alps,Austria)

The mollusc fauna of the early Middle Miocene (Langhian) intramontane Alpine Lake Groisenbach is described for the first time. The shells derive from the Feistring Formation in the Aflenz Basin in Austria, which was covered by Lake Groisenbach. The assemblage is moderately diverse with 12 gastropod and 2 bivalve species, suggesting shallow lacustrine and fluvial settings. Among the gastropods, only Theodoxus crenulatus (Klein, 1853) is known from other Miocene localities, whilst all other species are documented so far only from Lake Groisenbach. None of the Early and Middle Miocene lake systems of the Alpine-Carpathian Foredeep and the Balkan Peninsula displays any faunistic resemblance with this new fauna. Even coeval lake faunas from the close-by Graz Basin have no species in common with Lake Groisenbach. This pattern points to a surprising endemicity and biogeographic fragmentation in the Central European freshwater systems during the Early and Middle Miocene. The uniqueness of the newly described fauna is also indicated by the completely erratic occurrence of the otherwise African-Mediterranean genus Bulinus, which is unknown from all other central European Miocene freshwater systems. Emmericia roetzeli Harzhauser and Neubauer nov. sp., Nematurella zuschini Neubauer and Harzhauser nov. sp., Romania fastigata Neubauer and Harzhauser nov. sp., Odontohydrobia groisenbachensis Neubauer and Harzhauser nov. sp., Odontohydrobia pompatica Neubauer and Harzhauser nov. sp., Odontohydrobia styriaca Harzhauser and Neubauer nov. sp., Planorbis austroalpinus Harzhauser and Neubauer nov. sp., Gyraulus sachsenhoferi Harzhauser and Neubauer nov. sp., Bulinus corici Harzhauser and Neubauer nov. sp., Ferrissia crenellata Harzhauser and Neubauer nov. sp. and Stagnicola reinholdkunzi Harzhauser and Neubauer nov. sp. are introduced as new species.     

New data from Oman indicate benthic high biomass productivity coupled with low taxonomic diversity in the aftermath of the Permian–Triassic Boundary mass extinction

A new Early Triassic marine fauna is described from an exotic block (olistolith) from the Ad Daffah conglomerate in eastern Oman (Batain), which provides new insights into the ecology and diversity during the early aftermath of the Permian–Triassic Boundary mass extinction. Based on conodont quantitative biochronology, we assign a middle Griesbachian age to the upper part of this boulder. It was derived from an offshore seamount and yielded both nektonic and benthic faunas, including conodonts, ammonoids, gastropods and crinoid ossicles in mass abundance. This demonstrates that despite the stratigraphically near extinction at the Permian–Triassic Boundary, Crinoidea produced enough biomass to form crinoidal limestone as early as middle Griesbachian time. Baudicrinus, previously placed in Dadocrinidae, is now placed in Holocrinidae; therefore, Dadocrinidae are absent in the Early Triassic, and Holocrinidae remains the most basal crown‐group articulates, originating during the middle Griesbachian in the Tethyan Realm. Abundant gastropods assigned to Naticopsis reached a shell size larger than 20 mm and provide another example against any generalized Lilliput effect during the Griesbachian. Whereas the benthic biomass was as high as to allow the resumption of small carbonate factories, the taxonomic diversity of the benthos remained low compared to post‐Early Triassic times. This slow benthic taxonomic recovery is here attributed to low competition within impoverished post‐extinction faunas.     

Impact of fire and the recovery of molluscs in south‐east Australian salt marsh

Fire has long been recognised as a natural force in structuring Northern Hemisphere salt marshes, yet little is known about the impact of fire on molluscs and native vegetation dynamics of Southern Hemisphere coastal salt marshes. Following a fire at Ash Island, Hunter River New South Wales, Australia in the summer 2012, we assessed patterns of recovery through time of gastropod populations and resident salt marsh vegetation including biomass for three keystone native plant species, Native Rush (Juncus kraussii Hochst.), a chenopod (Sarcocornia quinqueflora Bunge ex Ungen‐Sternberg A.J. Scott), Salt Couch (Sporobolus virginicus, L. Kunth) and the invasive Spiny Rush (Juncus acutus). In temperate east‐coast Australian salt marshes, Spiny Rush is displacing native salt marsh vegetation. After twelve months, the biomass of Native Rush recovered to similar pre‐burn levels. While fire affected the abundance, richness and composition of the gastropod assemblage differences were also largely driven by spatial variability. Gastropod assemblages associated with two of the higher elevation native species (Native Rush and Salt Couch) were impacted the most by fire. Greater abundance (between 1 and 5 orders of magnitude difference in abundance) and richness of gastropods were found in unburnt compared with burnt Native Rush and Salt Couch vegetation, while more gastropods were found in Spiny Rush in one site. Species prevalent in burnt vegetation included larger species of gastropods Ophicardelus ornatus (Ferussac, 1821) and Phallomedusa solida (Martens, 1878) with an unexpected spike in number of the smaller gastropod Tatea huonensis (Tenison‐Woods, 1876) in the spiny rush at one site only. In salt marsh habitats, many gastropods have planktonic larval dispersal stages which are dependent on the tidal height for transport and the structural complexity provided by vegetation at settlement. Since fire appears to negatively affect salt marsh gastropod populations within structurally complex Native Rush and Salt Couch, due consideration of the importance of these refuges for gastropods is recommended when fire or other disturbances occur in ecologically endangered salt marsh in the Southern Hemisphere. Managers need to consider spatial heterogeneity of molluscs and their recovery in the event of fire in Southern Hemisphere salt marshes.     

The food and feeding habits of five freshwater and brackish-water fish species in Nigeria

The food and feeding habits of five economically important fresh-and brackish-water fishes, Channa obscura, Chrysichthys nigrodigitatus, Heterotis niloticus, Synodontis nigrita and Trachinotus maxillosus, were investigated. A number of techniques were used to carry out gut content analysis, including the Hynes 'point' system based on volume estimations expressed as relative percentages. Juveniles of C. nigrodigitatus were omnivorous, consuming 32% gastropods, 30% nematodes, 14% diatoms and 8% crustaceans, while adults were planktotrophic, consuming 23% diatoms, 33% Chlorophyceae and 22% crustaceans. Synodontis nigrita juveniles fed almost exclusively (91%) on nematodes, while adults were predominantly planktotrophic, their diet comprising 50% diatoms and 50% crustaceans. Trachinotus maxillosus was exclusively benthotrophic, consuming 99.5% gastropods and 0.46% nematodes. Heterotis niloticus was planktotrophic at the adult stage and consumed 72% crustaceans, 12% gastropods, 3% fish and about 90% planktotrophic at the juvenile phase. Chrysichthys obscura was purely benthotrophic at the juvenile stage, feeding 100% on nematodes, but fed mainly (89%) on fish at the adult stage.     

Palaeoenvironments of Early Miocene hominoid-bearing deposits at Napak,Uganda, based on terrestrial molluscs

Studies of fossil molluscs from Early Miocene deposits at Napak, Uganda, reveal that between 20 and 18.5 Ma, there were various kinds of palaeoenvironments on the flanks of the volcano ranging from forest to open country. The basal Iriri Member contains fluvial deposits from which freshwater gastropods and bivalves have been collected. The younger Napak Member is subaerial in facies, and has yielded a predominantly forest assemblage of gastropods, but with some indications of open country species at one of the sites. Napak has yielded a rich and diverse primate fauna, including the large ape Ugandapithecus major, the small ape Micropithecus clarki, the earliest known cercopithecid in the world (Prohylobates sp.) and several galagids (Mioeuoticus bishopi) among other less well-known taxa.     

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.