An evolutionary estuarine incursion: molecular differentiation and niche separation in Bornean Indothais snails (Rapaninae, Muricidae)

Ecological and biogeographic information for marine gastropods has lagged behind taxonomic, molecular and phylogenetic information, limiting understanding of evolutionary processes. We investigated niche separation and speciation in rapinine whelks that represent a rare deep estuarine penetration by muricid gastropods. We studied the genetic and ecological differentiation of four Indothais morphotypes, distributed along a salinity gradient in Brunei (Borneo, South East Asia). Our molecular analyses, based on three mitochondrial genes (COI, 12S rRNA and 16S rRNA) and one nuclear gene (28S rRNA), revealed three species having the following relationship [Indothais rufotincta (Tan & Sigurdsson, 1996), [I. javanica (Philippi, 1848), [I. gradata (Jonas, 1846)]]]. This pattern coincided with their salinity-related distributions, such that I. rufotincta occurred alone in the open coastal waters, and I. javanica and I. gradata extended into the upper estuary. This suggests speciation through divergent selection and local adaptation, specifically synapomorphic low salinity tolerance by I. javanica and I. gradata. The observed intertidal vertical separation where the species co-occur suggests competitive exclusion of the others by the most-recently evolved I. gradata. This species also showed greatest morphological and genetic variation, and unique niche expansion involving feeding on both hard surface and muddy sediment organisms. Our study presents a novel hypothesis for the speciation of these snails.

New taxa and new synonymy in Muricidae (Neogastropoda: Pagodulinae, Trophoninae, Ocenebrinae) from the Northeast Pacific

The results of an extensive examination of northeast Pacific muricid gastropods ranging from Aleutian Islands, Alaska, to mid-Baja California, is presented. Two new genera and 26 new species are described: In Pagodulinae: Abyssotrophon fusiformis n. sp., A. newmani n. sp., Boreotrophon cascadiensis n. sp., B. cordellensis n. sp., B. cortesianus n. sp., B. obesus n. sp., B. subapolyonis n. sp., B. vancouverensis n. sp., B. aleuticus n. sp., B. pseudotripherus n. sp., B. santarosensis n. sp., B. tannerensis n. sp. In Trophoninae: Warenia, n. gen., Scabrotrophon buldirensis n. sp, S. kantori n. sp., S. lima n. sp., S. macleani n. sp., S. moresbyensis n. sp., S. norafosterae n. sp., S. trifidus n. sp., Nipponotrophon exquisitus n. sp. In Ocenebrinae: Paciocinebrina n. gen., Nucella angustior n. sp., Paciocinebrina benitoensis n. sp., P. macleani n. sp., P. neobarbarensis n. sp., P. pseudomunda n. sp., P. thelmacrowae n. sp. New synonymy: Boreotrophon kamchatkanus Dall, 1902 (+ Trophonopsis nanus Ergorov, 1994); Paciocinebrina atropurpurea (Carpenter, 1865) (+ Tritonalia interfossa var. clathrata Dall, 1919, Ocinebra rubra Baker, 1891, Tritonalia tracheia Dall, 1919); P. barbarensis (Gabb, 1865) (+ Tritonalia interfossa var. beta Dall, 1919, Ocenebra keenae Bormann, 1946); P. circumtexta (Stearns, 1871) (+ Ocinebra circumtexta var. aurantia Stearns, 1895, Tritonalia circumtexta var. citrica Dall, 1919, Tritonalia lurida var. rotunda Dall, 1919); P. foveolata (Hinds, 1844) (+Tritonalia epiphanea Dall, 1919, Tritonalia fusconotata Dall, 1919); P. gracillima (Stearns, 1871) (+ Tritonalia gracillima var. obesa Dall, 1919, Ocinebra stearnsi Hemphill, 1911); P. interfossa (Carpenter, 1864) (+ Tritonalia interfossa alpha Dall, 1921); P. lurida (Middendorff, 1848) (+ Vitularia aspera Baird, 1863), P. sclera (Dall, 1919) (+ Coralliophila (Pseudomurex) kincaidi Dall, 1919). Generic assignments are changed for the following taxa: Boreotrophon kamchatkanus Dall, 1902; Warenia elegantula (Dall, 1907); Scabrotrophon stuarti (E.A. Smith, 1880); Paciocinebrina atropurpurea (Carpenter, 1865), P. barbarensis (Gabb, 1865), P. circumtexta (Stearns, 1871), P. crispatissima (Berry, 1953), P. foveolata (Hinds, 1844), P. fraseri (Oldroyd, 1920), P. gracillima (Stearns, 1871), P. grippi (Dall, 1911), P. interfossa (Carpenter, 1864), P. lurida (Middendorff, 1848), P. minor (Dall, 1919), P. munda (Carpenter, 1864), P. seftoni (Chace, 1958), P. sclera (Dall, 1919). Boreotrophon alborostratus Taki, 1938, is reinstated. Abyssotrophon Egorov, 1993 and Nodulotrophon Habe & Ito, 1965 are here assigned to Pagodulinae Barco et al., 2012, based on radula morphology.

Tetraclita (Cirripedia, Thoracica) tests as an important habitat for intertidal isopods and other marine and semi-terrestrial fauna on tropical rocky shores

The barnaclesTetraclita singaporensisandT. squamosahave a thick test to protect the animals against the diverse environmental stress of the tropical intertidal zone and also against predation by muricid gastropods. After the death of the barnacle, however, the empty test is often taken over by other marine fauna as well as semi-terrestrial animals. The sphaeromatid isopodDynamenella ptychurawas the most abundant inhabitant in empty tests observed in Singapore and Malaysia. Ovigerous crustaceans were common. Gastropod specimens comprised almost entirely juveniles of common intertidal species, including those of the littorinidsLittoraria articulataandL. strigata. Gastropod eggs and veligers, insect larvae, pupae and nymphs, and spider spiderlings and immatures were also present. The haminoeid gastropodSmaragdinellawas the dominant animal colonizing barnacles in a succession experiment. Temperature was significantly lower inside the empty test than outside, by 0.2°C. Almost 40% of the barnacle tests remained attached to the substratum for more than four months after the death of the individuals.

Mixed assemblages of drilling predators and the problem of identity in the fossil record: A case study using the muricid gastropod Ecphora

Drillholes made by naticid and muricid gastropods are frequently used in evolutionary and ecological studies because they provide direct, preservable evidence of predation. The muricid Ecphora is common in many Neogene Atlantic Coastal Plain assemblages in the United States, but is frequently ignored in studies of naticid predation. We used a combination of Pliocene fossil, modern beach, and experimentally derived samples to evaluate the hypothesis that Ecphora was an important source of drillholes in infaunal bivalve prey shared with naticids. We focused on the large, thick-shelled venerid, Mercenaria, which is commonly drilled by naticids today. Laboratory experiments, modern beach samples, and the published literature confirm that naticids preferentially drill near the umbo (significant clumping of holes), show a significant correlation between prey size and predator size (estimated by outer borehole diameter), and prefer Mercenaria <50 mm antero-posterior width when other prey are present. Fossil samples containing Ecphora (with or without other large muricids) show no drillhole site stereotypy (no significant clumping, greater variability in placement), no significant predator: prey size correlation, drilled prey shells larger than the largest modern naticids could produce in an experimental setting, and drillholes larger in diameter than those estimated for the largest Pliocene naticids, thus supporting our hypothesis. Substantial overlap in the placement of holes drilled by naticids and muricids, however, made identifying predators from drillhole position problematic. The lack of overlapping ranges of prey shell thickness between fossil and other samples precluded the use of drillhole morphology to establish predator identity (e.g., ratio of inner borehole diameter to outer borehole diameter, drillhole angle). Whereas the difficulty in determining predator identity from drillholes limits the types of analyses that can be reliably performed in mixed-predator assemblages, recognizing Ecphora as a prominent drilling predator creates the opportunity to investigate previously unrecognized questions.

Seasonal variations in the density of and corallivory by Drupella rugosa and Cronia margariticola (Caenogastropoda: Muricidae) from the coastal waters of Hong Kong: ‘plagues’ or ‘aggregations’?

Sixteen coral sites in the coastal waters of Hong Kong were examined for the corallivorous muricid gastropods Drupella rugosa and Cronia margariticola. These were recorded from all sites where there was significant hard coral cover and observed feeding upon species of Platygyra, Leptastrea, Stylocoeniella, Porites, Favites, Cyphastrea, Goniastrea, Favia, Acropora, Montipora, Pavona, Lithophyllon, Hydnophora, Echinophyllia and Plesiastrea. One large aggregation (~2000 individuals) of, mainly, D. rugosa was observed but much smaller groups (<20 individuals) were more typical. Five sites were chosen for more detailed study and surveyed during winter and summer. Despite being characterized by different coral communities, inter-site densities of D. rugosa were not significantly different and, usually, ~2 individuals · m2 were recorded. Seasonal differences were, however, significant with numbers greater during the summer, possibly related to reproduction. Feeding activity followed a similar pattern and was also largely confined to summer.Prey selection by Drupella rugosa was complex in the field and changed according to the relative abundance of each coral taxon. Acropora was strongly selected for at all sites where it was present and Montipora, Platygyra and Pavona were usually fed upon in greater proportions than their abundances. Leptastrea, Cyphastrea, Favites, Favia and Goniastrea were fed upon but in proportions lower than suggested by their abundances. Goniopora was never fed upon despite being relatively common. The seasonality of feeding, low density, rarity of large feeding aggregations, prey selection and aspects of the feeding behaviour, that is, generally only consuming the coral's coenenchyme (the polyps surviving), suggest that while D. rugosa is widespread in Hong Kong, and contrary to other views, it poses little, if any, threat to local coral communities. Thus, reported feeding clusters of D. rugosa are probably not ‘plague’ outbreaks but examples of seasonally fostered ‘aggregations’ of feeding (and probably reproducing) individuals. Indeed, no ‘plague-like’ outbreak of any species of Drupella has been reported upon in the literature since 1999.