Invasive and non-invasive lineages in Xenostrobus (Bivalvia: Mytilidae)

The mussel genus Xenostrobus comprises eight named extant species from the Indo-West Pacific, including Xenostrobus inconstans, Xenostrobus pulex and Xenostrobus securis from Australia, the latter two also occurring in New Zealand. Xenostrobus species are predominantly restricted to estuaries or sheltered marine habitats although X. pulex inhabits the rocky intertidal on open coasts. Xenostrobus has recently been suggested to be congeneric with the freshwater invasive species Limnoperna fortunei. Xenostrobus securis is itself invasive in the Far East and Europe. This study employed DNA sequences from cytochrome c oxidase (COI), the D1 expansion region of 28S rRNA and the internal transcribed spacers of the ribosomal cistrons to investigate species relationships and habitat transitions in Australasian Xenostrobus. It is unlikely that Xenostrobus and Limnoperna are congeneric as their COI sequences are very different. There was unexpected complexity in defining Xenostrobus species. Xenostrobus pulex from New Zealand is probably a distinct species to the Australian taxon of that name with the name X. neozelanicus available for that taxon. Xenostrobus inconstans and Australian X. pulex were not reciprocally monophyletic in COI analyses. The phylogeography of the COI haplotypes of X. securis suggests that it may be a cryptic species complex, although this possibility could not be confirmed by sequences of the other genes.     

The biology and functional anatomy of Modiolarca tumida (Musculus marmoratus) Bivalvia: Mytilidae

Modiolarca tumida (Hanley, 1843) is a member of the sub-family Crenellinae (Mytilidae). The preferred habitat of the species is the test of certain ascidians. The shell is dorsally flattened, which prevents it from cutting into the test during dorso-ventral contraction of the byssal retractors. The use of the byssus enables it to surround itself completely with host tissue. Adoption of the feeding posture involves the anterior-posterior contraction of the byssal retractors, which elevates the posterior margin above the host's surface using the anterior margin as the fulcrum against the host. Modiolarca tumida are attracted by the tunicin of the host, a process probably facilitated by the host's feeding currents. The smallest individuals are found round the oral aperture. Colonization of other parts of the host may result from surface migration as M. tumida can be highly mobile, crawling by means of the very extensible foot. It is during this process that individuals may be swept away in local currents and be forced to adopt a free-living existence.     

The effect of Lithophaga (Bivalvia: Mytilidae) boreholes on the strength of the coral Porites lobata

At Isla del Caño, Costa Rica, the main reef bioeroders are species of the boring bivalve, Lithophaga. These mussels are abundant in virtually all colonies of the main framework builder, Porites lobata. The bivalves have a significant effect on the strength of the colonies which, at a maximum compressive and bending strength of 20 and 5 MN/m², respectively, are already the lowest ever measured for corals. Presence of bivalve boreholes is inversely logarithmically associated with coral strength, with the greatest decrease in strength occurring with the first boreholes.     

Mitochondrial DNA polymorphisms and sperm motility in Mytilus edulis (Bivalvia: Mytilidae)

The system of mitochondrial DNA (mtDNA) inheritance in Mytilus and other bivalves, termed doubly uniparental inheritance (DUI), is novel among animals. Males pass on their male transmitted (M-type) mtDNA from fathers to their sons whereas females pass on their female transmitted (F-type) mtDNA from mothers to both sons and daughters. Thus, Mytilus males contain two distinct types of mtDNA. Interestingly, sperm contains only the paternal mtDNA. Phylogenetic analysis has shown that some female types have been able to switch their route of inheritance. These "recently masculinized" mitochondrial genomes behave as a typical M-type in that they are transmitted from generation to generation through sperm. Because the "recently masculinized" and "standard" male mitotypes in M. edulis exhibit approximately 8.7% amino acid sequence divergence, we hypothesized that these differences could affect mitochondrial, and hence sperm, functions. Furthermore, since recently masculinized mitotypes have been shown to replace standard male types periodically over evolutionary timescales, we tested the hypothesis that sperm swimming speeds would be greater for males with recently masculinized M-type genomes. Sperm activity was videotaped, digitized and tracked. A linear mixed effects model found no significant difference in linear velocities or curvilinear speeds between the mitotypes suggesting that swimming speeds are similar for both in the period shortly after spawning.     

Larval development of the invasive charru mussel,Mytella strigata (Bivalvia: Mytilidae)

ABSTRACT

The South American charru mussel, Mytella strigata, was recently recorded in Singapore waters, possibly introduced into Southeast Asia through shipping. The mussels have rapidly spread across estuarine coastal mudflats. Adult mussels were collected, spawned in aquaria and larvae were successfully cultured to the juvenile stage in the laboratory. The larval morphology and development of M. strigata is described in this paper. D-shaped veligers were produced within 20 h of fertilization and were approximately 75 µm in shell length. These larvae were capable of settlement two weeks post fertilization. Given an adequate amount of food, they were able to grow up to 1 mm in shell length within 30 days. The larval shell of M. strigata possesses anterodorsal G2 hinge teeth as distinct wavy ledges, with a pitted resilial ridge clearly evident in the juvenile shell.     

Size and age composition of populations and growth of Mytilus trossulus (Bivalvia: Mytilidae) in the subtidal area of western Kamchatka

Fairly large Mytilus trossulus beds were found in the subtidal zone at depths to 40 m off the western coast of Kamchatka. The size and age composition of local populations and growth rates of M. trossulus in this region and at Atlasova Island were studied. Significant variability of the shell shape of mollusks was found. In the western Kamchatka subtidal, M. trossulus grew faster than in the northern Kuril Islands and eastern Kamchatka. Mussels attained commercial size (50 mm in shell length) in the fourth year of life in the former case and in the fifth-sixth year in other cases. Mollusks of different ages, from 1 to 8–9 year olds, were present in western Kamchatka populations; i.e., juvenile recruitment takes place regularly, while growth rates vary. Growth and population structure of M. trossulus are discussed in relation to the habitat conditions on the western Kamchatka shelf.     

Distribution pattern, population structure, and growth of Septifer bilocularis (Bivalvia: Mytilidae) on reefs of Southern Vietnam

The features of the spatial distribution, size and age structure of populations, and growth of the bivalve mollusk Septifer bilocularis (Linnaeus, 1758) were investigated for the first time in the waters of Vietnam. It is shown that in the subtidal area, on reefs well protected from wave effects, S. bilocularis occupies a corallogenous substrate constructed of dead branchy colonies of Acropora and Porites and forms settlements there with a population density and biomass exceeding those in populations located along the open coast and developing on massive coral colonies of the genera Porites, Favia, and Favites. The extensive accumulation of sediments in the inner part of Vanphong Bay (Khanh Hoa Province) correlated with a decrease in the size and age ranges for populations of S. bilocularis. It is found that in places with regularly active hydrodynamics the mollusks grew somewhat more slowly and formed a more convex shell than on reefs protected from surf. The longevity of S. bilocularis estimated both from the greatest observed age and from the growth equation of Bertalanffy was somewhat greater in the former case compared with the latter one, but in general did not exceed 11 years for both cases. The results of the studies are discussed from the position of morphophysiological adaptations of sessile bivalve mollusks to life in contrasting environmental conditions of the upper subtidal zone.     

Cytogenetic characterization of the invasive mussel species Xenostrobus securis Lmk. (Bivalvia: Mytilidae)

The chromosomes of the invasive black-pigmy mussel (Xenostrobus securis (Lmk. 1819)) were analyzed by means of 4',6-diamidino-2-phenylindole (DAPI) / propidium iodide (PI) and chromomycin A3 (CMA) / DAPI fluorescence staining and fluorescent in situ hybridization using major rDNA, 5S rDNA, core histone genes, linker histone genes, and telomeric sequences as probes. The diploid chromosome number in this species is 2n = 30. The karyotype is composed of seven metacentric, one meta/submetacentric, and seven submetacentric chromosome pairs. Telomeric sequences appear at both ends of every single chromosome. Major rDNA clusters appear near the centromeres on chromosome pairs 1 and 3 and are associated with bright CMA fluorescence and dull DAPI fluorescence. This species shows five 5S rDNA clusters close to the centromeres on four chromosome pairs (2, 5, 6, and 8). Three of the four core histone gene clusters map to centromeric positions on chromosome pairs 7, 10, and 13. The fourth core histone gene cluster occupies a terminal position on chromosome pair 8, also bearing a 5S rDNA cluster. The two linker histone gene clusters are close to the centromeres on chromosome pairs 12 and 14. Therefore, the use of these probes allows the unequivocal identification of 11 of the 15 chromosome pairs that compose the karyotype of X. securis.     

Functional phenotypic plasticity of the endolithic mytilid Leiosolenus curtus (Lischke, 1874) (Bivalvia: Mytilidae)

In Sagami Bay, Japan, Leiosolenus curtus (Lischke, 1874) bores into dead shells of rock barnacles (Tetraclita japonica) in Manazuru, calcareous sand-rock in Oiso, weathered tuff in Miura and agglomerates formed by the tubes of clam worms (Spirobranchus kraussii) in Kanaya. In order to understand the relationships between morphology and environment, the shell morphologies and genetic characteristics of L. curtus, together with the calcium content of the different bored substrata, were analysed. Individuals from Kanaya and Manazuru were characterised by a chalk-like encrustation and a more rounded and inflated shell shape, while those from Oiso and Miura had a striated encrustation. The molecular phylogenetic tree and haplotype network showed that the genetic sequences of the individuals from all investigated localities were very similar, and implied that they belong to the same species. The striated encrustation of the individuals from Oiso and Miura presumably enables them to scrape those residues not dissolved by the mucus secreted by the mantle gland, and thereby to bore into non-calcareous substrata. The more rounded and inflated shell shape of the individuals from Manazuru is thought to be advantageous for boring into thin substrata.     

Genetic and morphometric characterization of mussels (Bivalvia: Mytilidae) from mid-atlantic hydrothermal vents.

Mussels were collected from deep-sea hydrothermal vents along the Mid-Atlantic Ridge. Specimens from the Snake Pit site were previously identified genetically and anatomically as Bathymodiolus puteoserpentis, but the relationships of mussels from other sites (Logatchev and Lucky Strike) were unclear. Molecular genetic and morphological techniques were used to assess differences among these mussel populations. The results indicate that the range for B. puteoserpentis extends from Snake Pit to Logatchev, and that an unnamed second species, B. n. sp., occurs at Lucky Strike. Analysis of mitochondrial NADH dehydrogenase subunit 4 (ND4) revealed 13% sequence divergence between the two species. Nei's genetic distance (D) based on 14 allozyme loci was 0.112. A multivariate morphometric analysis yielded a canonical discriminant function that correctly identified individuals from these sites to species 95% of the time.     

Phylogenetic evidence for role-reversals of gender-associated mitochondrial DNA in Mytilus (Bivalvia: Mytilidae)

Distinct gender-associated mitochondrial DNA (mtDNA) lineages (i.e., lineages which are transmitted either through males or through females) have been demonstrated in two families of bivalves, the Mytilidae (marine mussels) and the Unionidae (freshwater mussels), which have been separated for more than 400 Myr. The mode of transmission of these M (for male-transmitted) and F (for female-transmitted) molecules has been referred to as doubly uniparental inheritance (DUI), in contrast to standard maternal inheritance (SMI), which is the norm in animals. A previous study suggested that at least three origins of DUI are required to explain the phylogenetic pattern of M and F lineages in freshwater and marine mussels. Here we present phylogenetic evidence based on partial sequences of the cytochrome c oxidase subunit I gene and the 16S RNA gene that indicates the DUI is a dynamic phenomenon. Specifically, we demonstrate that F lineages in three species of Mytilus mussels, M. edulis, M. trossulus, and M. californianus, have spawned separate lineages which are now associated only with males. This process is referred to as "masculinization" of F mtDNA. By extension, we propose that DUI may be a primitive bivalve character and that periodic masculinization events combined with extinction of previously existing M types effectively reset the time of divergence between conspecific gender-associated mtDNA lineages.      

A mitochondrial phylogeography of Brachidontes variabilis (Bivalvia: Mytilidae) reveals three cryptic species

This study examined genetic variation across the range of Brachidontes variabilis to produce a molecular phylogeography. Neighbour joining (NJ), minimum evolution (ME) and maximum parsimony (MP) trees based on partial mitochondrial DNA sequences of 16S-rDNA and cytochrome oxidase (COI) genes revealed three monophyletic clades: (1) Brachidontes pharaonis s.l. from the Mediterranean Sea and the Red Sea; (2) B. variabilis from the Indian Ocean; (3) B. variabilis from the western Pacific Ocean. Although the three clades have never been differentiated by malacologists employing conventional morphological keys, they should be ascribed to the taxonomic rank of species. The nucleotide divergences between Brachidontes lineages (between 10.3% and 23.2%) were substantially higher than the divergence between congeneric Mytilus species (2.3–6.7%) and corresponded to interspecific divergences found in other bivalvia, indicating that they should be considered three different species. Analysis of the 16S-rDNA sequences revealed heteroplasmy, indicating dual uniparental inheritance (DUI) of mtDNA in the species of Brachidontes collected in the Indian Ocean, but not in the species in the Pacific nor in the species in the Red Sea and the Mediterranean Sea. When we employed the conventional estimate of the rate of mitochondrial sequence divergence (2% per million years), the divergence times for the three monophyletic lineages were 6–11 Myr for the Indian Ocean and Pacific Ocean Brachidontes sp. and 6.5–9 Myr for the Red Sea and Indian Ocean Brachidontes sp . Thus, these species diverged from one another during the Miocene (23.8–5.3 Myr). We infer that a common ancestor of the three Brachidontes species probably had an Indo-Pacific distribution and that vicariance events, linked to Pleistocene glaciations first and then to the opening of the Red Sea, produced three monophyletic lineages.     

Genetics and taxonomy of Chilean smooth-shelled mussels, Mytilus spp. (Bivalvia: Mytilidae)

It has been previously established that native smooth-shelled mussels in southern South America possess close evolutionary affinities with Northern-Hemisphere Mytilus edulis L. 1758 (McDonald et al. (1991) [5]). This result has since been challenged by authors claiming that Chilean mussels should be considered a local subspecies of M. galloprovincialis Lmk. 1819. Moreover, morphological, physiological, ecotoxicological and molecular genetic studies on Chilean smooth-shelled mussels still frequently refer to ‘M. chilensis’ Hupé 1854, even though the previous discovery of alien M. galloprovincialis and considerable heterogeneity in shell morphology among samples collected along the Chilean shores raise concerns that different Mytilus spp. species might have been included under ‘M. chilensis’. Here we reviewed the molecular and morphological data available on smooth-shelled mussels from Chile in an attempt to clarify both their genetic composition and their taxonomic status. Using multivariate analysis on sample × allozyme-frequency matrices, we confirmed the widespread occurrence of the Southern-Hemisphere form of M. edulis along the shores from the North Patagonia region of Chile to the southern tip of the South American continent. The populations sampled in southern central Chile showed some evidence of slight introgression from Southern-Hemisphere M. galloprovincialis. Morphological characterization of a sample from Dichato in southern central Chile was consistent with its previous genetic identification as Mediterranean M. galloprovincialis. The occurrence of Southern-Hemisphere M. galloprovincialis in Punta Arenas at the southern tip of the South American continent was also reported. Southern-Hemisphere M. edulis, including native Chilean smooth-shelled Mytilus, should be assigned subspecific rank and named M. edulis platensis d’Orbigny 1846.     

A hybrid zone between hydrothermal vent mussels (Bivalvia: Mytilidae) from the Mid-Atlantic Ridge

This study provides the first example of a hybrid zone between animal taxa distributed along the mid-ocean ridge system. We examined the distribution and genetic structure of deep-sea hydrothermal vent mussels (Bivalvia: Mytilidae) along a 2888-km portion of the Mid-Atlantic Ridge between 37 degrees 50' N and 14 degrees 45' N latitude. Mitochondrial DNA (mtDNA), allozymes and multivariate-morphometric evidence discriminated between individuals of a northern species, Bathymodiolus azoricus, and a southern species, B. puteoserpentis, that were separated by an intermediate ridge segment almost devoid of mussels. A small sample of mussels from Broken Spur, a vent locality along this intermediate zone, revealed a mixed population with gene frequencies and morphology that were broadly intermediate to those of the northern and southern species. Multilocus clines in mtDNA and allozyme frequencies were centred over the intermediate zone. We consider intrinsic and extrinsic processes that might limit genetic exchange across this hybrid zone.     

Parasitism in species of Bathymodiolus (Bivalvia: Mytilidae) mussels from deep-sea seep and hydrothermal vents

Bivalve species, especially mussels, are biomass dominants in many deep-sea chemosynthetic ecosystems. As in shallow-water environments, parasites are likely to be important factors in the population dynamics of bivalve communities in chemosynthetic ecosystems, but there has been little study of parasitism in deep-sea seep or vent molluscs. In this study, parasite types, diversity, prevalence, infection density and non-infectious indicators of stress or disease as related to host age, reproductive condition, and endosymbiont density were assessed in mussels (Bathymodiolus heckerae) from 2 seep sites and mussels (B. puteoserpentis) from 2 vent sites. We identified 10 microbial or parasitic agents in histological sections. Parasite types included 3 viral-like gut inclusions, 2 rickettsia-like gill inclusions, a rickettsia-like mantle inclusion, a bacterial gill-rosette, a chlamydia-like gut inclusion, gill-dwelling ciliates, and an unidentified inclusion in gut tissues. Parasite species richness was greater in seep mussels than in vent mussels, with the seep mussels possessing 9 types of parasites compared to 2 in the vent mussels. One of the viral-like inclusions infecting the seep mussel B. heckerae was pathogenic, causing lysis of the digestive tubules. The prevalence and intensity of infection by this pathogen were greater in hosts with shell lengths less than 100 mm. Mussels from all 4 sites also exhibited intense infiltration of tissues and blood spaces by enlarged hemocytes. Hemocytic infiltration (hemocytosis) showed variable degrees of severity that were not associated with other host factors examined.     

The trematode invasion and accumulation on heavy metals by the mollusc Colletopterum ponderosum (Bivalvia: Unionidae: Anodontinae)

The influence of the trematode invasion on the accumulation of heavy metals (Cu, Zn, Cd, Pb) by various organs of the mollusc Colletopterum ponderosum have been investigated. An increase of the coefficients of accumulation and bottom biological accumulation, and also changes in the accumulation in pairs of heavy metals were noted in the infected molluscs.