Marine Biodiversity

Flyingfish (Exocoetidae) species diversity and habitats in the eastern tropical Pacific Ocean
Lewallen EA, van Wijnen AJ, Bonin CA and Lovejoy NR
Flyingfishes are large enough to eat zooplankton, small enough to be consumed by top predators, and therefore form a central mid-trophic component of tropical epipelagic marine food webs. Characterizing patterns of flyingfish abundance, distribution, and habitat preference have important implications for understanding both localized and generalized functions of marine ecosystems. The eastern tropical Pacific Ocean (ETP) supports many flyingfish species and their predators, yet no studies to date have identified oceanographic factors that define flyingfish habitats or estimate species richness and diversity at broad taxonomic and geographic scales. In this study, we analyzed 11,125 flyingfish representing 25 species and all 7 named genera, collected from the ETP over a 21-year period. We applied spatially-explicit analysis methods (ARCGIS, DIVA-GIS, MAXENT) and compared specimen locality data to remotely-sensed oceanographic data, and previously described oceanographic partitions. Our results show that is the most abundant genus (49%), and the most abundant species (32%) of flyingfishes in the ETP. Mean sea surface temperature was most important for defining flyingfish habitats (19.2-41.7%) and species richness (highest in the North Equatorial Current). Additionally, flyingfish species diversity was found to be highest in coastal regions of the study area (Shannon indices > 1.5). Together, these results provide unprecedented characterizations of a mid-trophic epipelagic community in an economically valuable region during a time when sea surface temperatures are predicted to increase as a result of global climate change.
Isolation and characterisation of polymorphic microsatellite loci for studies of the big blue octopus,
McKeown NJ, Taylor AL and Shaw PW
The big blue octopus, , occurs on coral reefs throughout the Indo-Pacific region from East Africa to the Hawaiian Islands, wherein it is of great ecological and socio-economic importance. However, many components of its intraspecific biodiversity, such as population structure, are unresolved due to a lack of informative genetic markers. To address this issue, which may compromise conservation and sustainability efforts, the development and characterisation of the first species-specific microsatellite loci for are described here. The eight loci were characterised by the genotyping of 40 adults from Madagascar, which revealed an average of 13.5 alleles per locus (range 9-18). The observed and expected heterozygosity per locus ranged from 0.432 to 0.949 and from 0.481 to 0.989, respectively. No evidence of linkage disequilibrium was detected between pairs of loci. Genotype proportions at six loci conformed to Hardy-Weinberg equilibrium expectations, with two loci exhibiting significant heterozygote deficits. These loci are applicable to multiple areas of eco-evolutionary research and, thus, represent a valuable resource for future studies of .
Infestation of shore crab gills by a free-living mussel species
Poulter R, Oliver PG, Hauton C, Sanders T and Ciotti BJ
Parasitic and commensal species can impact the structure and function of ecological communities and are typically highly specialized to overcome host defences. Here, we report multiple instances of a normally free-living species, the blue mussel Linnaeus, 1758, inhabiting the branchial chamber of the shore crab (Linnaeus, 1758) collected from widely separated geographical locations. A total of 127 were examined from four locations in the English Channel, one location in the Irish Sea and two locations at the entrance of the Baltic Sea. The branchial chambers of three crabs (one from the English Channel and two from Gullmar Fjord, Sweden) were infested with mussels resembling the genus . Sequencing at the Me15/16 locus on the polyphenolic adhesive protein gene confirmed the identity as . Bivalve infestation always occurred in larger red male individuals. Up to 16 mussels, ranging from 2 to 11 mm in shell length, were found in each individual, either wedged between gill lamellae or attached to the branchial chamber inner wall. This is one of the first reports of a bivalve inhabiting crustacean gills and is an intriguing case of a normally free-living prey species infesting its predator.
Mitochondrial genome and polymorphic microsatellite markers from the abyssal sponge Lim & Wiklund, 2017: tools for understanding the impact of deep-sea mining
Taboada S, Kenny NJ, Riesgo A, Wiklund H, Paterson GLJ, Dahlgren TG and Glover AG
The abyssal demosponge is endemic to the Clarion - Clipperton Zone (CCZ) in the NE Pacific, a region with abundant seafloor polymetallic nodules and of potential interest for mining. encrusts on these nodules and is an abundant component of the ecosystem. To assess the impact of mining operations, it is crucial to understand the genetics of this species, because its genetic diversity and connectivity across the area may be representative of other nodule-encrusting invertebrate epifauna. Here we describe and characterize 14 polymorphic microsatellite markers from this keystone species using Illumina MiSeq, tested for 75 individuals from three different areas across the CCZ, including an Area of Particular Environmental Interest (APEI-6) and two areas within the adjacent UK1 mining exploration area. The number of alleles per locus ranged from 3 to 30 (13.33 average alleles for all loci across areas). Observed and expected heterozygosity ranged from 0.909-0.048 and from 0.954-0.255, respectively. Several loci displayed significant deviation from the Hardy-Weinberg equilibrium, which appears to be common in other sponge studies. The microsatellite loci described here will be used to assess the genetic structure and connectivity on populations of the sponge across the CCZ, which will be invaluable for monitoring the impact of mining operations on its habitat. Also, we provide the annotated mitochondrial genome of , compare its arrangement with other closely related species, and discuss the phylogenetic framework for the sponge after Maximum Likelihood and Bayesian Inference analyses using nucleotide and amino acid sequences data sets separately.
Watersiporidae (Bryozoa) in Iberian waters: an update on alien and native species
Reverter-Gil O and Souto J
Species of the genus comprise an important invasive fouling group but are difficult to identify up to species level. This problem, in conjunction with the recent re-description of several member species, requires the revision of previous records and newly collected material in order to more precisely determine their actual presence and distribution. Here, we revise the identity and distribution of alien and native species of Watersiporidae in Iberian waters based on newly collected material, historical collections, and bibliographic data. Four species of are now known from here. is the only native species, present in the Spanish Mediterranean. seems to have been introduced relatively recently in Iberian and European Atlantic waters and has been expanding to other Atlantic localities. was collected for the first time in Europe at the SW Spanish Atlantic coast in 1990 and recently in Mediterranean marinas. is known in Iberian waters from two localities in the Gulf of Cadiz and in Gibraltar. With the recent redescription of the genus , is no longer a member. A new watersiporid genus, gen. nov., is described for this species. In Iberian waters, comb. nov. is considered a native species, frequent and abundant in shallow waters along the Atlantic coast.
Dismantling the (Busk, 1852) species complex (Bryozoa, Cheilostomata): two new species from European waters
Souto J, Nascimento KB, Reverter-Gil O and Vieira LM
New research on bryozoans has determined that formerly widespread species are in many cases complexes of similar, but distinct, species with more restricted distributions. Notwithstanding, the limits of distribution are still unresolved for many taxa, and occasionally a wide distribution is confirmed. has been considered a widespread species, distributed throughout the Southern Hemisphere, parts of northern Pacific and Atlantic Oceans and the Mediterranean Sea. This study examines the Magellanic-type material, together with other historic samples and new specimens collected in the western Mediterranean and Adriatic, and for the first time, presents specimens from the European North Atlantic. Morphological comparisons and biometric analysis show the existence of three different species among the specimens studied. A redescription of based on the type specimen is presented, and two new species are described: sp. nov. from the Northeast Atlantic and sp. nov. from the Mediterranean Sea. These results indicate that s.l. is a large complex of species and that most specimens from different parts of the world must be revised.
sp. nov. (Crustacea: Amphipoda) exhibits ontogenetic vertical stratification across abyssal and hadal depths in the Atacama Trench, eastern South Pacific Ocean
Weston JNJ, Espinosa-Leal L, Wainwright JA, Stewart ECD, González CE, Linley TD, Reid WDK, Hidalgo P, Oliva ME, Ulloa O, Wenzhöfer F, Glud RN, Escribano R and Jamieson AJ
S.I. Smith in Scudder, 1882 (Crustacea: Amphipoda) are prevalent scavengers of the benthopelagic community from bathyal to hadal depths. While a well-studied genus, molecular systematic studies have uncovered cryptic speciation and multiple undescribed lineages. Here, we apply an integrative taxonomic approach and describe the tenth species, sp. nov., based on specimens from the 2018 Atacamex and RV Sonne SO261 Expeditions to the southern sector of the Peru-Chile Trench, the Atacama Trench (24-⁠21°S). sp. nov. is a large species, max. observed length 83.2 mm, possesses diagnostic features, including a short gnathopod 1 palm and a chelate gnathopod 2 palm, and a distinct genetic lineage based on a 16S rRNA and COI phylogeny. This species is a dominant bait-attending fauna with an extensive bathymetric range, spanning from 4974 to 8081 m. The RV Sonne SO261 specimens were recovered along a 10-station transect from abyssal to hadal depths and further examined for demographic and bathymetric-related patterns. Ontogenetic vertical stratification was evident across the trench axis, with only juveniles present at abyssal depths (4974-6025 m). Total length-depth analysis revealed that the size of females was unrelated to depth, whereas juveniles followed a sigmoidal relationship with a step-up in size at depths >7200 m. Thus, these bathymetric trends suggest that juveniles and females employ differing ecological strategies in subduction trench environments. This study highlights that even dominant and ecologically important species are still being discovered within the abyssal and hadal environments. Continued systematic expeditions will lead to an improved understanding of the eco-evolutionary drivers of speciation in the world's largest ecosystem.
Hydrothermal vent fauna of the Galápagos Rift: updated species list with new records
Chen C, Jamieson JW and Tunnicliffe V
The sighting of giant bivalves and tubeworms at the Rose Garden vent field on the Galápagos Rift in 1977 marked the discovery of hydrothermal vents, a turning point for modern biology. The following decade saw a flurry of taxonomic descriptions of vent endemic species from the first vents. With the finding of high-temperature "black smokers" on the East Pacific Rise, exploration shifted away from Galápagos. A faunal list of Galápagos vents with 65 species was published in 1991, then updated to 74 species in 2006. Since then, few expeditions returned to the Galápagos Rift. Here, we revisited several Galápagos vents including recently confirmed high-temperature sites and inactive sulfide mounds. From our collecting efforts and observations, as well as revisions from the literature, we update the faunal list to 92 species including 15 new records, restricted to obvious vent associates. Accurate regional faunal lists are important for understanding the biogeography of vent fauna, and our list will also be valuable for setting management strategies.
Appreciation to reviewers
Bryozoan diversity on a whale bone: an uncommon substrate from the continental shelf off NW Spain
Souto J and Reverter-Gil O
Carcasses and bones of whales are interesting yet poorly understood habitats for invertebrates and microbial communities. Nevertheless, bryozoans have never before been reported in this particular habitat at the present day. Here, we present the first study on a bryozoan community colonizing a whale bone from the continental shelf off NW Spain at 128 m depth. Thirty-three species of bryozoans were identified, representing 50% of the known bryozoan diversity from this area and depth range. Despite this high biodiversity, no potential whale-fall specialists were detected: all the species are already known in the area, with several being more frequent in shallower or deeper waters. Our results support the hypothesis that the number of specialist taxa on whale-fall habitats decreases drastically on whale remains at depths of less than 260 m. These results also support the theory that whale-falls can serve as intermediate habitats where no other suitable substrate is present, facilitating habitat colonization.
Benthic megafaunal biodiversity of the Charlie-Gibbs fracture zone: spatial variation, potential drivers, and conservation status
Keogh P, Command RJ, Edinger E, Georgiopoulou A and Robert K
The Charlie-Gibbs Fracture Zone (CGFZ) is a prominent geological feature offsetting the Mid-Atlantic Ridge (MAR), consisting of two parallel fractures, creating a highly variable seafloor bathymetry. It has been defined as the most important latitudinal biodiversity transitional zone on the MAR. Despite this recognition, the faunal communities living on the fracture zone have not been extensively described. A remotely operated vehicle (ROV) was utilised during the TOSCA (Tectonic Ocean Spreading at the Charlie-Gibbs Fracture Zone) survey. The survey included five ROV video transects at depths between 560 and 2900 m. The objectives of the study were to use this video footage to quantify benthic megafaunal density and biodiversity patterns on the CGFZ and their environmental drivers. Species accumulation curves and generalised additive modelling show that depth and the presence of hard substrates play an important role in explaining species richness and abundance at the CGFZ. Coral taxa showed highest abundance between depths of 1500 and 2000 m, while sponge taxa were more abundant between 1750 and 2250 m. A dense sponge aggregation was identified on a ridge feature at 2250 m depth. The high biodiversity and presence of dense sponge aggregations and coral gardens found in this study highlight the need for detailed surveys to help support decisions made by governing bodies on the protection status of the CGFZ.
The molluscan assemblage of a pristine meadow in the eastern Mediterranean
Holzknecht M and Albano PG
The seagrass forms extensive meadows in the Mediterranean Sea. Studies on their associated highly diverse invertebrate assemblages are limited to the western Mediterranean. The eastern Mediterranean, however, is a basin undergoing rapid change due to the synergistic effects of climate warming, biological invasions and other human stressors that are driving native biodiversity to regional-scale collapses. We here surveyed the shelled molluscan assemblage of a meadow in Plakias, south-western Crete, the first such study in the eastern Mediterranean Sea. This area has increased its yearly mean temperature by 1 °C in the last 20 years and is under heavy pressure by Lessepsian species. We sampled across a 5- to 20-m depth gradient, in two seasons to capture intra-annual variation and the leaf and rhizome strata separately. Against our expectations, the molluscan assemblage proved to be highly diverse, with species richness, dominant species and trophic guilds comparable to healthy western Mediterranean ones, and with a negligible non-indigenous component. The diversity of the native community (following the biotic resistance hypothesis) and oxygen supersaturation in the meadow may cause greater resistance to biological invasions and warming, respectively, suggesting that meadows may act as a precious refugium for native biodiversity in the fast changing eastern Mediterranean Sea.
New species and new records of bryozoan species from fouling communities in the Madeira Archipelago (NE Atlantic)
Souto J, Ramalhosa P, Ferrario J, Png-Gonzalez L, Álvarez S, Gestoso I, Nogueira N and Canning-Clode J
Hull fouling is considered to be the most significant vector of introduction of marine non-indigenous species (NIS) in the Madeira Archipelago (NE Atlantic) because these islands provide a vital passage route for many ships. The transfer of species between boat hulls and artificial substrates in marinas is known to be high. Bryozoans are among the most common groups of marine invertebrates growing on this type of substrate. In recent years, significant advances have been made in our knowledge about the biodiversity of bryozoans in the Madeira Archipelago. Nonetheless, the currently recognized numbers remain far from reflecting the actual bryozoan species richness. In this context, we examine bryozoan samples stemming from NIS monitoring surveys on artificial substrates along the southern coast of the Madeira Archipelago, in four recreational marinas and in two offshore aquaculture farms. This has yielded new information about ten bryozoan species. Two of them, sp. nov. and sp. nov., are described for the first time, although at least the first one was previously recorded from Madeira but misidentified. , , , and are recorded for the first time in Madeira. Moreover, the material of was compared with the type, and a biometric analysis was performed with material from the Atlantic and Mediterranean. All samples identified as in both regions are the same species, and the variations described in the literature apparently reflect high intracolonial variability. Finally, we provide new information for the descriptions of 4 additional bryozoans, namely, sp. aff, , , and .
Wave-swept coralliths of Saba Bank, Dutch Caribbean
Hoeksema BW, Hassell D, Meesters EHWG and van Duyl FC
During a recent reef coral survey at the submarine Saba Bank (Eastern Caribbean), an uncommon and diverse assemblage of unattached scleractinian corals (coralliths) was encountered, which has not been reported from the Atlantic before. Four different types of these free-living (unattached) corals were distinguished. They were observed on a relatively flat seafloor (15-20 m deep) with poor coral cover and full exposure to oceanic swell. Much of the substratum was not consolidated and consisted mainly of sand and fragments of branching coralline algae. One of the four types is the (1) anthocyathus stage in the life history of the free-living species and . The other three are coralliths formed as ecophenotypic varieties: (2) spheroidal-amoeboidal (= globular and (sub)massive) in , , , and ; (3) tumbleweed-like (= globular and ramose) in and ; and (4) discoidal (flat and circular with short branches) in and possibly in cf. . This assemblage of free-living corals is likely related to a combination of abiotic factors consisting of wave exposure (swell), depths that waves can reach, a horizontal sea floor with little relief, an unconsolidated substratum, and low coral cover.
nom. nov. (Echinodermata, Holothuroidea) from bathyal bottoms of Galicia (NW Iberian Peninsula)
Urgorri V, Candás M, Díaz-Agras G, Cunha-Veira X, Gómez-Rodríguez C and Míguez-Rodríguez L
It is proposed to name the Atlantic holothuria Cherbonnier, 1969, whose denomination is already occupied by (Semper, 1867) from the Philippines, as nom. nov. 843 specimens of nom. nov., collected between 417 and 1191 m deep in the NW and W of Galicia and in the Galicia Bank, were studied. A detailed description of the external and internal anatomy of nom. nov. is made by studying the ossicles and the skeletal structure by means of scanning electron microscopy (SEM) as well as the introvert, calcareous ring, retractor muscles, water-vascular system, digestive system, respiratory trees, and reproductive system by means of micro-computed tomography (micro-CT). The habitat, feeding system, and geographical distribution of nom. nov. are also described.
Abundance and morphology of , observed at the Clarion-Clipperton Zone
Durden JM, Simon-Lledo E, Gooday AJ and Jones DOB
is an important trace fossil characterised by a regular hexagonal structure and typical of ancient deep-ocean habitats as far back as the Ordovician. It is represented in modern deep-sea settings by , known from the Mid-Atlantic Ridge, the South Atlantic, and off eastern Australia. Here we report the occurrence of in the Clarion Clipperton Zone (CCZ), abyssal equatorial Pacific, an area characterised by polymetallic nodule fields. At the study site within the International Seabed Authority northeastern Area of Particular Environmental Interest (APEI-6), appeared as a compact, regular pattern of small circular openings on the seafloor, each pattern interpreted as reflecting the activity of an individual organism. The patterns had a mean size (maximum dimension) of 45 mm ± 16 mm SD (n = 841) and occurred at a density of 0.33 individuals m. Most (82%) were interrupted by nodules, but those that were not displayed both regular (59%) and irregular (41%) forms, the former having equal numbers of rows along the three axes (6 x 6 x 6 and 8 x 8 x 8). In both size and morphology, our traces were more similar to the Australian examples than to those from the Mid-Atlantic Ridge.
Non-indigenous species of Bryozoa from anthropogenic habitats in the Bay of Cádiz (South Iberian Peninsula)
Souto J and Reverter-Gil O
Samples of bryozoans collected from two localities in the Bay of Cádiz with different levels of anthropogenic impact are studied. A total of 25 species was identified, of which 8 are considered non-indigenous (NIS), 12 as native and 5 cryptogenic. A new species of , sp. nov., is here described, and corrections to the diagnosis of the genus are proposed. The species appears to be a recent immigrant in Cádiz, although it could also be present in the eastern Mediterranean. and the genus itself are recorded for the first time in the eastern Atlantic. is recorded for the first time in Spanish waters. was already recorded in the Iberian Peninsula, but previous records should be confirmed. Other species, such as , , and , were already recorded in the Bay of Cádiz very recently.