The Chinese pond mussel Sinanodonta woodiana demographically outperforms European native mussels.

Unionid mussels are essential for the integrity of freshwater ecosystems but show rapid worldwide declines. The large-sized, thermophilic Chinese pond mussel Sinanodonta woodiana s.l., however, is a successful global invader, spread with commercially traded fish encysted with mussel larvae; its negative impacts on native mussels are expected. Here, we exploit a natural experiment provided by a simultaneous introduction of S. woodiana and four species of native unionids for water filtration to a pond in north-eastern Poland. Sinanodonta woodiana established a self-sustaining population and persisted for 19 years in suboptimal thermal conditions (mean annual temperature, 7.4 °C; mean temperature of the coldest month, - 3.7 °C, 73-day mean yearly ice-formation), extending the known limits of its cold tolerance. Over four study years, its frequency increased, and it showed higher potential for population growth than the native mussels, indicating possible future dominance shifts. Outbreaks of such sleeper populations are likely to be triggered by increasing temperatures. Additionally, our study documents the broad tolerance of S. woodiana concerning bottom sediments. It also points to the importance of intentional introductions of adult individuals and the bridgehead effect facilitating its further spread. We argue that S. woodiana should be urgently included in invasive species monitoring and management programmes.

Global biogeography of chemosynthetic symbionts reveals both localized and globally distributed symbiont groups.

In the ocean, most hosts acquire their symbionts from the environment. Due to the immense spatial scales involved, our understanding of the biogeography of hosts and symbionts in marine systems is patchy, although this knowledge is essential for understanding fundamental aspects of symbiosis such as host-symbiont specificity and evolution. Lucinidae is the most species-rich and widely distributed family of marine bivalves hosting autotrophic bacterial endosymbionts. Previous molecular surveys identified location-specific symbiont types that "promiscuously" form associations with multiple divergent cooccurring host species. This flexibility of host-microbe pairings is thought to underpin their global success, as it allows hosts to form associations with locally adapted symbionts. We used metagenomics to investigate the biodiversity, functional variability, and genetic exchange among the endosymbionts of 12 lucinid host species from across the globe. We report a cosmopolitan symbiont species, Candidatus Thiodiazotropha taylori, associated with multiple lucinid host species. Ca. T. taylori has achieved more success at dispersal and establishing symbioses with lucinids than any other symbiont described thus far. This discovery challenges our understanding of symbiont dispersal and location-specific colonization and suggests both symbiont and host flexibility underpin the ecological and evolutionary success of the lucinid symbiosis.

Molecular Characterization of Complement Component 3 (C3) in the Pearl Oyster Pinctada fucata Improves Our Understanding of the Primitive Complement System in Bivalve.

As the central component in the complement system, complement component 3 (C3) plays essential roles in both the innate and adaptive immune responses. Here, a C3 gene (designated as pf-C3) was obtained from the pearl oyster Pinctada fucata by RT-PCR and rapid amplification of cDNA ends (RACE). The pf-C3 cDNA consists of 5,634 bp with an open reading frame (ORF) of 5,193 bp encoding a protein of 1,730 amino acids with a 19 residue signal peptide. The deduced pf-C3 protein possessed the characteristic structural features present in its homologs and contained the A2M_N_2, ANATO, A2M, A2M_comp, A2M_recep, and C345C domains, as well as the C3 convertase cleavage site, thioester motif, and conserved Cys, His, and Glu residues. Phylogenetic analysis revealed that pf-C3 is closely related to the C3s from other mollusks. Pf-C3 mRNA was expressed in all examined tissues including gill, digestive gland, adductor muscle, mantle and foot, while the highest expression was found in the digestive gland. Following the challenge with Vibrio alginolyticus, pf-C3 expression was significantly induced in hemocytes. Luciferase reporter assays indicated that pf-C3a could activate the NF-κB signal pathway in HEK293T cells. Further knockdown of pf-C3 by specific siRNA could significantly reduce the phagocytosis of V. alginolyticus by hemocytes in vitro. These results would help increase understanding of the function of C3 in the invertebrate immune system and therefore provide new insights into the roles of the primitive complement system in invertebrates.

Systematic redescription of Solen/ (Ensisolen/) tehuelchus/ and Ensis/ macha/ (Bivalvia: Solenoidea) from Argentina, southwestern Atlantic Ocean.

In the present study, bivalves belonging to the superfamily Solenoidea that inhabit the Argentinean Sea are redescribed. The specimens studied in this work were collected by scuba diving in the Gulf of San José, Argentina. Additional specimens from different malacological collections were revised to update the geographical distribution of valid species. According to the literature, there exist two species for the "Argentinean" and "Magellan" biogeographical provinces. However, ten nominal taxa, with uncertain taxonomic status, have been mentioned for the study area. Field work, local collection survey and type material examination allow confirming two valid species: Solen (Ensisolen) tehuelchus Hanley (Solenidae) and Ensis macha (Molina) (Pharidae). A morphological comparison between S. tehuelchus and S. gaudichaudi Chenu and S. obliqua Spengler is provided as well type locality, repository and synonymy list. The type material of all related species are illustrated whenever possible. Establishing the valid names of these species is the first step towards more integrative studies including genetic and zoogeographical analyses.

Phylogenetic Relationships and Adaptation in Deep-Sea Mussels: Insights from Mitochondrial Genomes.

Mitochondrial genomes (mitogenomes) are an excellent source of information for phylogenetic and evolutionary studies, but their application in marine invertebrates is limited. In the present study, we utilized mitogenomes to elucidate the phylogeny and environmental adaptation in deep-sea mussels (Mytilidae: Bathymodiolinae). We sequenced and assembled seven bathymodioline mitogenomes. A phylogenetic analysis integrating the seven newly assembled and six previously reported bathymodioline mitogenomes revealed that these bathymodiolines are divided into three well-supported clades represented by five Gigantidas species, six Bathymodiolus species, and two "Bathymodiolus" species, respectively. A Common interval Rearrangement Explorer (CREx) analysis revealed a gene order rearrangement in bathymodiolines that is distinct from that in other shallow-water mytilids. The CREx analysis also suggested that reversal, transposition, and tandem duplications with subsequent random gene loss (TDRL) may have been responsible for the evolution of mitochondrial gene orders in bathymodiolines. Moreover, a comparison of the mitogenomes of shallow-water and deep-sea mussels revealed that the latter lineage has experienced relaxed purifying selection, but 16 residues of the atp6, nad4, nad2, cob, nad5, and cox2 genes have underwent positive selection. Overall, this study provides new insights into the phylogenetic relationships and mitogenomic adaptations of deep-sea mussels.

A High-Quality Reference Genome for a Parasitic Bivalve with Doubly Uniparental Inheritance (Bivalvia: Unionida).

From a genomics perspective, bivalves (Mollusca: Bivalvia) have been poorly explored with the exception for those of high economic value. The bivalve order Unionida, or freshwater mussels, has been of interest in recent genomic studies due to their unique mitochondrial biology and peculiar life cycle. However, genomic studies have been hindered by the lack of a high-quality reference genome. Here, I present a genome assembly of Potamilus streckersoni using Pacific Bioscience single-molecule real-time long reads and 10X Genomics-linked read sequencing. Further, I use RNA sequencing from multiple tissue types and life stages to annotate the reference genome. The final assembly was far superior to any previously published freshwater mussel genome and was represented by 2,368 scaffolds (2,472 contigs) and 1,776,755,624 bp, with a scaffold N50 of 2,051,244 bp. A high proportion of the assembly was comprised of repetitive elements (51.03%), aligning with genomic characteristics of other bivalves. The functional annotation returned 52,407 gene models (41,065 protein, 11,342 tRNAs), which was concordant with the estimated number of genes in other freshwater mussel species. This genetic resource, along with future studies developing high-quality genome assemblies and annotations, will be integral toward unraveling the genomic bases of ecologically and evolutionarily important traits in this hyper-diverse group.

Natural hybridization between pen shell species: Pinna rudis and the critically endangered Pinna nobilis may explain parasite resistance in P. nobilis.

Recently, Pinna nobilis pen shells population in Mediterranean Sea has plummeted due to a Mass Mortality Event caused by an haplosporidian parasite. In consequence, this bivalve species has been included in the IUCN Red List as "Critically Endangered". In the current scenario, several works are in progress to protect P. nobilis from extinction, being identification of hybrids (P. nobilis x P. rudis) among survivors extremely important for the conservation of the species.Morphological characteristics and molecular analyses were used to identify putative hybrids. A total of 10 individuals of each species (P. nobilis and P. rudis) and 3 doubtful individuals were considered in this study. The putative hybrids showed shell morphology and mantle coloration intermingled exhibiting both P. nobilis and P. rudis traits. Moreover, the analyses of 1150 bp of the 28S gene showed 9 diagnostic sites between P. rudis and P. nobilis, whereas hybrids showed both parental diagnostic alleles at the diagnostic loci. Regarding the multilocus genotypes from the 8 microsatellite markers, the segregation of two Pinna species was clearly detected on the PCoA plot and the 3 hybrids showed intermediate positions.This is the first study evidencing the existence of hybrids P. nobilis x P. rudis, providing molecular methodology for a proper identification of new hybrids. Further studies testing systematically all parasite-resisting isolated P. nobilis should be undertaken to determine if the resistance is resulting from introgression of P. rudis into P. nobilis genome and identifying aspects related to resistance.

Host Diversity Affects Parasite Diversity: A Case Study Involving Unionicola spp. Inhabiting Freshwater Mussels.

Water mites of the genus Unionicola are common parasites of freshwater mussels, living on the gills or mantle of their hosts and using these tissues as sites of oviposition. Although surveys of this mite fauna among North American unionid mussels indicate that these mites represent highly diverse assemblages, we know very little regarding the determinants of Unionicola species diversity among their molluscan hosts. The present study addresses the relationship between host diversity and mite diversity for Unionicola assemblages associated with unionid mussels of North America. The results of this study found a significantly positive relationship between host species richness and mite species richness, adding to a growing body of evidence that host diversity is an important determinant of parasite diversity. In recent years, molecular sequence data have discovered cryptic biodiversity among unionid mussels, yielding revisions in the nomenclature and systematic taxonomy of the group. DNA sequence variation has also revealed cryptic species complexes among Unionicola mites. Collectively, these findings suggest that the results of the present study may be underestimating species richness among mites and their host mussels. Unfortunately, human perturbations are known to have caused high recent rates of extinction in the mussel and mite faunas of North America and could play a major role in influencing patterns of species richness for this host-parasite system moving forward.

Comparative mitogenomic analysis of the superfamily Tellinoidea (Mollusca: Bivalvia): Insights into the evolution of the gene rearrangements.

The superfamily Tellinoidea is widespread and contains approximately 180 living species, which is one of the most diverse and representative groups among the bivalves. In order to extend our knowledge on evolution of tellinoidean species, we newly determined five tellinoidean mitochondrial genomes (mitogenomes). The newly determined mitogenome vary in size from 16,333 to 16,986 bp. The results show that the genome size and genome organization are conserved in tellinoideans. However, gene arrangement and the location of the major non-coding region (NCR) show diversity. The atp8 gene presents in all the five new mitogenomes. Two trnK and trnP genes were detected in Gari togata mitogenome. Phylogenetic analysis supports the monophyly of Tellinoidea, however, it's family Psammobiidae is polyphyletic. CREx analysis suggests that the gene order of Nuttallia olivacea is assumed as the most primitive condition of Tellinoidea. We map the gene order onto the phylogeny and infers the possible gene rearrangement scenarios among tellinoidean mitogenomes. The mitochondrial gene rearrangement is a useful information that help reassessing the phylogeny of Tellinoidea. Phylogenetic relationship and gene arrangement analyses suggest that a careful review for the current taxonomy of the family Psammobiidae is required.

Genome-wide identification, characterisation and expression analysis of the ALAS gene in the Yesso scallop (Patinopecten yessoensis) with different shell colours.

Porphyrins, one of the most common shell pigments, are by-products of the haem pathway. 5-Aminolaevulinate synthase (ALAS) is the first and rate-limiting enzyme in this pathway and has been well studied in vertebrate species. However, the function of ALAS in shell colouration has been poorly studied in molluscs, which are renowned for their colourful shells. In the present study, an ALAS gene, named PyALAS, was identified through whole-genome scanning in the Yesso scallop (Patinopecten yessoensis), an economically and evolutionarily important bivalve species in which the shell colour represents polymorphism. Two conserved domains were detected in the PyALAS protein sequence, including a Preseq-ALAS domain and a 5-ALAS domain, confirming the identification of PyALAS. Phylogenetic analysis of the ALAS proteins among various invertebrate and vertebrate species revealed a high consistency between the molecular evolution of ALAS and the species taxonomy. PyALAS was ubiquitously expressed in most adult tissues of the Yesso scallop. The left mantle expressed a significantly higher level of PyALAS than the right side in brown scallops, whereas there was no significant difference in white scallops. Significantly different expression levels of PyALAS was also detected between the two different shell colour strains. These data indicate that PyALAS plays an important role in shell colouration in Yesso scallops and the present study provides new insights into the molecular mechanism of shell colouration in molluscs.

New freshwater mussel taxa discoveries clarify biogeographic division of Southeast Asia.

While a growing body of modern phylogenetic research reveals that the Western Indochina represents a separate biogeographic subregion having a largely endemic freshwater fauna, the boundaries of this subregion are still unclear. We use freshwater mussels (Unionidae) as a model to reconstruct spatial patterns of freshwater biogeographic divides throughout Asia. Here, we present an updated freshwater biogeographic division of mainland Southeast Asia and describe 12 species and 4 genera of freshwater mussels new to science. We show that the Isthmus of Kra represents a significant southern biogeographic barrier between freshwater mussel faunas of the Western Indochina and Sundaland subregions, while the Indian and Western Indochina subregions are separated by the Naga Hills, Chin Hills, and Rakhine Yoma mountain ranges. Our findings highlight that the freshwater bivalve fauna of Southeast Asia primarily originated within three evolutionary hotspots (Western Indochina, Sundaland, and East Asian) supplemented by ancient immigrants from the Indian Subcontinent.

Massive bioconstructions built by Neopycnodonte cochlear (Mollusca, Bivalvia) in a mesophotic environment in the central Mediterranean Sea.

The present paper provides a multidisciplinary fine-scale description of a Mediterranean mesophotic new habitat dominated by the bivalve Neopycnodonte cochlear (Poli, 1795), building large and thick pinnacles on vertical cliffs at two study areas along the southern Italian coast. The pinnacles, constituted by a multilayered aggregation of living and dead specimens of N. cochlear, were interconnected with each other to form a framework of high structural complexity, never observed before for this species. The bioconstruction, considerably extended, resulted very complex and diversified in the associated community of structuring organisms. This latter included 165 taxa attributable to different ecological groups occurring in different microhabitats of the bioconstruction. Among the secondary structuring taxa there were scleractinians, serpulids and bryozoans, all contributing to the deposition of calcium carbonate, and poriferans, helping to bind shells together or eroding carbonate by boring species. In comparison with coralligenous sensu stricto and the recently described Mediterranean mesophotic coral reef, the Neopycnodonte bioconstruction showed peculiar features, since it lacked the major contribution of encrusting coralline algae and scleractinians as reef builders, respectively.

Embryo and larval development of the yellow clam Mesodesma mactroides (Reeve, 1854) (Mesodesmatidae) in laboratory.

The yellow clam Mesodesma mactroides (Reeve, 1854) is a sand mollusc with historical and socioeconomic importance in Brazil, Uruguay and Argentina. A guaranteed form to access a successful reestablishment of the species in their natural environment is directly linked to their reproduction biology. Then, our report introduces the embryonic and larval development of the yellow clam reared in laboratory for such purposes. M. mactroides broodstock were selected as specimens who possess a mean total shell length and weight of 66 ± 3.82 mm and 27.15 ± 4.07 g for an afterwards spawn induction through stripping technique. Regarding the embryonic development, newly fertilized oocytes exhibited a mean diameter of 51.20 ± 6.64 µm. The first polar corpuscle, trochophores and D-veliger appeared at 20 min, 18 and 24 h after fertilization, respectively. Umbonate and pediveliger larvae were noticed on the 8th and 25th day, respectively, with complete metamorphosis occurring only at the 27th day, when all larvae were retained in a 200 µm nylon mesh. Therefore, with that basic understanding of the embryonic and larval development of M. mactroides in the laboratory, forwards studies will focus in establish a technological package for this species.

First mussel settlement observed in Antarctica reveals the potential for future invasions.

Global biodiversity is both declining and being redistributed in response to multiple drivers characterizing the Anthropocene, including synergies between biological invasions and climate change. The Antarctic marine benthos may constitute the last biogeographic realm where barriers (oceanographic currents, climatic gradients) have not yet been broken. Here we report the successful settlement of a cohort of Mytilus cf. platensis in a shallow subtidal habitat of the South Shetland Islands in 2019, which demonstrates the ability of this species to complete its early life stages in this extreme environment. Genetic analyses and shipping records show that this observation is consistent with the dominant vectors and pathways linking southern Patagonia with the Antarctic Peninsula and demonstrates the potential for impending invasions of Antarctic ecosystems.

Integrative taxonomy, biogeography and conservation of freshwater mussels (Unionidae) in Russia.

Freshwater mussels are ecosystem engineers and keystone species in aquatic environments. Unfortunately, due to dramatic declines this fauna is among the most threatened globally. Here, we clarify the taxonomy and biogeography of Russian Unionidae species based on the most comprehensive multi-locus dataset sampled to date. We revise the distribution and assess the conservation status for each species. This fauna comprises 16 native species from 11 genera and 4 tribes: Anodonta, Pseudanodonta (Anodontini); Amuranodonta, Beringiana, Buldowskia, Cristaria, Sinanodonta (Cristariini); Middendorffinaia, Nodularia, Unio (Unionini); and Lanceolaria (Lanceolariini). No country-level endemic species are known in Russia, except for Buldowskia suifunica that may also occur in China. Sinanodonta woodiana, a non-native species, was introduced from China. Russia comprises the northern parts of Western and Eastern Palearctic subregions. The first subregion with six species encompasses a huge area from the western boundary of Russia to the Lena Basin in Siberia. The second subregion with 10 species covers the Amur Basin, rivers east of the Lena Basin, coastal basins of the Japan Sea, and the North Pacific Islands. The fauna of Russia primarily includes widespread generalist species that are here considered Least Concern (LC). However, Buldowskia suifunica and Sinanodonta lauta have restricted distributions and are assessed here as Vulnerable (VU) and Endangered (EN), respectively.

Comparative phylogenomics reveal complex evolution of life history strategies in a clade of bivalves with parasitic larvae (Bivalvia: Unionoida: Ambleminae).

Freshwater mussels are a species-rich group with biodiversity patterns strongly shaped by a life history strategy that includes an obligate parasitic larval stage. In this study, we set out to reconstruct the life history evolution and systematics in a clade of freshwater mussels adapted to parasitizing a molluscivorous host fish. Anchored hybrid enrichment and ancestral character reconstruction revealed a complex pattern of life history evolution with host switching and multiple instances of convergence, including reduction in size of larvae, increased fecundity, and growth during encapsulation. Our phylogenomic analyses also recovered non-monophyly of taxa exhibiting multiple traits used as the basis for previous taxonomic hypotheses. Taxa with axe-head shaped glochidia were resolved as paraphyletic, but our results strongly suggest the complex morphology is an adaptation to reduce larval size, with reduction in size further accentuated in taxa previously assigned to Leptodea. To more accurately reflect the evolutionary history of this group, we make multiple systematic changes, including the description of a new genus, Atlanticoncha gen. nov., and the synonymy of the genus Leptodea under Potamilus. Our findings contribute to the growing body of literature showing that cladistic hypotheses based solely on morphological characters, including larval morphology, can be flawed in freshwater mussels.

Strong genetic differentiation of the clam Meretrix lamarckii in the China Sea revealed by mitochondrial DNA marker.

The hard clam Meretrix lamarckii is ecologically and economically important in the coastal regions of China. In this study, we evaluated the genetic diversity and population structure of six M. lamarckii populations in the East China Sea and the South China Sea using mitochondrial cytochrome c oxidase subunit 1 (COI) and cytochrome b (Cytb) genes. We obtained 582 bp of partly sequences comprising 28 novel haplotypes of COI gene from 138 specimens and 1168 bp of partly sequences comprising 22 novel haplotypes of Cytb gene from 125 specimens. The haplotype diversity of COI and Cytb genes ranged from 0.606 to 0.862 and 0.562 to 0.863, respectively. The nucleotide diversity ranged from 0.0015 to 0.0038 in COI gene and ranged from 0.0007 to 0.0032 in Cytb gene. Thus, there is moderate-level genetic diversity in M. lamarckii in the China Sea. The F-statistics showed that the Zhoushan (ZS) and Xiangshan (XS) populations were significantly (P < 0.01) differed from the populations of Wenzhou (WZ), Zhangpu (ZP), Shantou (ST), and Zhanjiang (ZJ) in both COI and Cytb genes. Both haplotypes network and plot of STRUCTURE analysis suggested obviously genetic divergence between East China Sea and South China Sea regions. Knowledge on genetic variation and population structure of M. lamarckii populations along the Southeast China Sea obtained from this study will support the aquaculture management and conservation of M. lamarckii in China.

Polymorphisms of LAP3 gene and their association with the growth traits in the razor clam Sinonovacula constricta.

Leucine aminopeptidase 3 (LAP3) is an important proteolytic enzyme that catalyzes the hydrolysis of leucine residues from the amino termini of protein or peptide substrates and plays a critical role in protein metabolism and growth. In the present study, we investigated the full-length cDNA sequence of the LAP3 gene in Sinonovacula constricta (ScLAP3) using expressed sequence tags and rapid amplification of cDNA ends. The full-length ScLAP3 cDNA was 2885 bp, with a 1560 bp open reading frame encoding 519 amino acids. Sequence analysis revealed that ScLAP3 shared 70.9% identity with LAP3 from the blood clam Tegillarca granosa and 62.0-68.0% with other species. ScLAP3 was expressed in all six tested tissues, with significantly higher expression levels in the foot compared with mantle, adductor muscle, liver, gills, and siphon tissues in adults (P < 0.01). In the eight developmental stages, ScLAP3 expression gradually increased, with significantly higher levels in D-shaped larvae compared with other developmental stages (P < 0.01), suggesting that it may be involved in the formation of certain organs during early development. Association analysis identified three shared single nucleotide polymorphisms (SNPs), c.1073A > G, c.1139C > T and c.1154A > G in exons of ScLAP3 gene from 177 individuals of two groups, one selective strain and one wild population, which had significant effects on growth traits of S. constricta. The results provided candidate genetic markers to assist selective breeding of razor clams toward improved growth.

Paleoecological insights from fossil freshwater mollusks of the Kanapoi Formation (Omo-Turkana Basin, Kenya).

The Early Pliocene Kanapoi Formation of the Omo-Turkana Basin consists of two fluvial/deltaic sedimentary sequences with an intermediate lacustrine sequence that was deposited in Paleolake Lonyumun, the earliest large lake in the basin. Overall, the geology and vertebrate paleontology of the Kanapoi Formation are well studied, but its freshwater mollusks, despite being a major component of the benthic ecosystem, have not been subjected to in-depth study. Here I present the first treatment of these mollusks, which have been retrieved mainly from the lacustrine but also from the upper fluvial sediments, with a focus on paleoecological implications. Overall, the freshwater mollusk fauna is reasonably diverse and contains the gastropods Bellamya (Viviparidae), Melanoides (Thiaridae), Cleopatra (Paludomidae) and Gabbiella (Bithyniidae), as well as the unionoid bivalves Coelatura, Pseudobovaria (Unionidae), Aspatharia, Iridina (Iridinidae) and Etheria (Etheriidae). Material is typically recrystallized and lithified and its taphonomy suggests deposition in a system with intermediate energy, such as a beach, with post-depositional deformation and abrasion. The mollusk assemblage is indicative of perennial, fresh and well-oxygenated waters in the Kanapoi region. It suggests that Paleolake Lonyumun had largely open shores with limited vegetation and that swampy or ephemeral backwaters were rare. Overall, these findings support earlier paleoecological interpretations based on the fish assemblage of Paleolake Lonyumun at Kanapoi. Moreover, mollusk assemblages from this lake are very similar across the Omo-Turkana Basin (Nachukui, Usno, Mursi and Koobi Fora Formations) suggesting that the lacustrine paleoecological conditions found in the Kanapoi Formation existed throughout the basin.

Mechanics unlocks the morphogenetic puzzle of interlocking bivalved shells.

Brachiopods and mollusks are 2 shell-bearing phyla that diverged from a common shell-less ancestor more than 540 million years ago. Brachiopods and bivalve mollusks have also convergently evolved a bivalved shell that displays an apparently mundane, yet striking feature from a developmental point of view: When the shell is closed, the 2 valve edges meet each other in a commissure that forms a continuum with no gaps or overlaps despite the fact that each valve, secreted by 2 mantle lobes, may present antisymmetric ornamental patterns of varying regularity and size. Interlocking is maintained throughout the entirety of development, even when the shell edge exhibits significant irregularity due to injury or other environmental influences, which suggests a dynamic physical process of pattern formation that cannot be genetically specified. Here, we derive a mathematical framework, based on the physics of shell growth, to explain how this interlocking pattern is created and regulated by mechanical instabilities. By close consideration of the geometry and mechanics of 2 lobes of the mantle, constrained both by the rigid shell that they secrete and by each other, we uncover the mechanistic basis for the interlocking pattern. Our modeling framework recovers and explains a large diversity of shell forms and highlights how parametric variations in the growth process result in morphological variation. Beyond the basic interlocking mechanism, we also consider the intricate and striking multiscale-patterned edge in certain brachiopods. We show that this pattern can be explained as a secondary instability that matches morphological trends and data.