Microbial signatures of protected and impacted Northern Caribbean reefs: changes from Cuba to the Florida Keys.

There are a few baseline reef-systems available for understanding the microbiology of healthy coral reefs and their surrounding seawater. Here, we examined the seawater microbial ecology of 25 Northern Caribbean reefs varying in human impact and protection in Cuba and the Florida Keys, USA, by measuring nutrient concentrations, microbial abundances, and respiration rates as well as sequencing bacterial and archaeal amplicons and community functional genes. Overall, seawater microbial composition and biogeochemistry were influenced by reef location and hydrogeography. Seawater from the highly protected 'crown jewel' offshore reefs in Jardines de la Reina, Cuba had low concentrations of nutrients and organic carbon, abundant Prochlorococcus, and high microbial community alpha diversity. Seawater from the less protected system of Los Canarreos, Cuba had elevated microbial community beta-diversity whereas waters from the most impacted nearshore reefs in the Florida Keys contained high organic carbon and nitrogen concentrations and potential microbial functions characteristic of microbialized reefs. Each reef system had distinct microbial signatures and within this context, we propose that the protection and offshore nature of Jardines de la Reina may preserve the oligotrophic paradigm and the metabolic dependence of the community on primary production by picocyanobacteria.

Development of a Handheld Submersible Chemiluminescent Sensor: Quantification of Superoxide at Coral Surfaces.

Reactive oxygen species (ROS) are produced via various photochemical, abiotic, and biological pathways. The low concentration and short lifetime of the ROS superoxide (O2•-) make it challenging to measure in natural systems. Here, we designed, developed, and validated a DIver-operated Submersible Chemiluminescent sensOr (DISCO), the first handheld submersible chemiluminescent sensor. The fluidic system inside DISCO is controlled by two high-precision pumps that introduce sample water and analytical reagents into a mixing cell. The resultant chemiluminescent signal is quantified by a photomultiplier tube, recorded by a miniature onboard computer and monitored in real time via a handheld underwater LED interface. Components are contained within a pressure-bearing housing (max depth 30 m), and an external battery pack supplies power. Laboratory calibrations with filtered seawater verified instrument stability and precision. Field deployment in Cuban coral reefs quantified background seawater-normalized extracellular superoxide concentrations near coral surfaces (0-173 nM) that varied distinctly with coral species. Observations were consistent with previous similar measurements from aquaria and shallow reefs using a standard benchtop system. In situ quantification of superoxide associated with corals was enabled by DISCO, demonstrating the potential application to other shallow water ecosystems and chemical species.

Two-year temporal response of benthic macrofauna and sediments to hypoxia in a tropical semi-enclosed bay (Cienfuegos, Cuba).

Hypoxia is the depletion of dissolved oxygen below 2 mg O(2)/L. Relatively few studies on hypoxia and its effects on benthic macrofauna have been done in tropical marine ecosystems. This study describes the temporal response of the water column, sediments and macrofauna to seasonal hypoxia in a semi-enclosed bay (Cienfuegos, Caribbean Sea). The Calisito site was sampled monthly from June 2010 until February 2012, yielding 21 sampling times. At each sampling event water and sediment samples were collected for measuring the abiotic variables (temperature, salinity, dissolved oxygen, nutrients, redox potential discontinuity, silt/clay and organic matter content) and macrofauna (abundance and species richness). Temperature and surface salinity followed a typical temporal pattern during the summer/rainy and the winter/dry periods. Salinity stratification occurred in the rainy period, lasting three months in 2010 and six months in 2011. The bottom water dissolved oxygen indicated hypoxic and anoxic events during the wet periods of 2010 and 2011 associated with salinity stratification, low hydrodynamics and oxidation of the accumulated organic matter. Over the study period, 817 individuals were collected and identified. Polychaetes were the dominant group in terms of abundance (57 % of total) followed by mollusks (41%). Hypoxia (and occasionally anoxia) caused strong deleterious effects on the abundance and species richness of macrofaunal communities in the study site. The most abundant polychaetes were opportunistic species with high tolerance to hypoxic conditions: Prionospio steenstrupi, Polydora sp.and Paraprionospio pinnata. Most of them colonized relatively fast once hypoxia ended. Persistent species such as Caecum pulchellum and Parvanachis obesa were present during hypoxia with fluctuating densities and apparently recover to higher abundances when normoxic conditions are re-established. Macoma tenta and Tellina consobrina colonized approximately 1-2 months later than the first polychaete peak during normoxia. Probably, the deleterious effects of hypoxia on the macrofauna were intensified by negative interspecific relationships such as competition by suitable space and predation. The recolonization of macrofauna depended possibly on local transport by currents within the bay because the connection with the Caribbean Sea is relatively limited. In summary, seasonal hypoxia in Cienfuegos Bay influences the water and sediment geochemistry and reduces both the abundance and diversity of macrofauna.

[Variations of the infracommuity during the ontogeny of Chromis cyanea (Perciformes: Pomcentride) in the North coast of Havana, Cuba]. / Variaciones de la infracomunidad parásita durante la ontogenia de Chromis cyanea (Perciformes: Pomacentridae) en la costa norte de La Habana, Cuba.

The infracommunty or parasites or unromis cyanea (Pisces: Pomacentridae) was studied along the ontogenetic development in the North coast of Havana, Cuba. The objectives were: a- to prove that the core species appears before the strange and stochastic species and they are responsible for the structure in the infracommunity, b- to determine if there is a relationship among the ecological describers of the parasitic infracommunity with the total length. A total of 278 specimens of C. cyanea were captured during the dry season (April) of 2010. The body size range was from 1.5 to 11.5 cm including juvenile and adults. We collected 2 861 parasite specimens belonging to 20 taxa: Crustacea (5), Nematoda (5), Trematoda (4), Cestoda (2), Monogenea (2), Turbellaria (1) and Ciliophora (1). The taxa Tetraphyllidea and Anilocra chromis constituted the core of the parasitic infracommunity. The sequence of appearance and persistence of these taxa during the ontogenetic development, supported the hypothesis of the core species. The changes in the infracommunity, from 6 cm body size, could be the result of an accumulative effect combined with changes of the diet that caused the ingestion of new parasite infective stages. We concluded that the ontogenetic development of C. cyanea constitutes an important variable in the formation of the parasitic infracommunity.

Taxonomic, phylogenetic, and functional diversity of mollusk death assemblages in coral reef and seagrass sediments from two shallow gulfs in Western Cuban Archipelago.

Mollusk death assemblages are formed by shell remnants deposited in the surficial mixed layer of the seabed. Diversity patterns in tropical marine habitats still are understudied; therefore, we aimed to investigate the taxonomic, phylogenetic, and functional diversity of mollusk death assemblages at regional and local scales in coral reef sands and seagrass meadows. We collected sediment samples at 11 sites within two shallow gulfs in the Northwestern Caribbean Sea and Southeastern Gulf of Mexico. All the shells were counted and identified to species level and classified into biological traits. We identified 7113 individuals belonging to 393 species (290 gastropods, 94 bivalves, and nine scaphopods). Diversity and assemblage structure showed many similarities between gulfs given their geological and biogeographical commonalities. Reef sands had higher richness than seagrasses likely because of a more favorable balance productivity-disturbance. Reef sands were dominated by epifaunal herbivores likely feeding on microphytobenthos and bysally attached bivalves adapted to intense hydrodynamic regime. In seagrass meadows, suspension feeders dominated in exposed sites and chemosynthetic infaunal bivalves dominated where oxygen replenishment was limited. Time averaging of death assemblages was likely in the order of 100 years, with stronger effects in reef sands compared to seagrass meadows. Our research provides evidence of the high taxonomic, phylogenetic, and functional diversity of mollusk death assemblages in tropical coastal sediments as result of the influence of scale-related processes and habitat type. Our study highlights the convenience of including phylogenetic and functional traits, as well as dead shells, for a more complete assessment of mollusk biodiversity.

Circumtropical distribution and cryptic species of the meiofaunal enteropneust Meioglossus (Harrimaniidae, Hemichordata).

Hemichordata has always played a central role in evolutionary studies of Chordata due to their close phylogenetic affinity and shared morphological characteristics. Hemichordates had no meiofaunal representatives until the surprising discovery of a microscopic, paedomorphic enteropneust Meioglossus psammophilus (Harrimaniidae, Hemichordata) from the Caribbean in 2012. No additional species have been described since, questioning the broader distribution and significance of this genus. However, being less than a millimeter long and superficially resembling an early juvenile acorn worm, Meioglossus may easily be overlooked in both macrofauna and meiofauna surveys. We here present the discovery of 11 additional populations of Meioglossus from shallow subtropical and tropical coralline sands of the Caribbean Sea, Red Sea, Indian Ocean, and East China Sea. These geographically separated populations show identical morphology but differ genetically. Our phylogenetic reconstructions include four gene markers and support the monophyly of Meioglossus. Species delineation analyses revealed eight new cryptic species, which we herein describe using DNA taxonomy. This study reveals a broad circumtropical distribution, supporting the validity and ecological importance of this enigmatic meiobenthic genus. The high cryptic diversity and apparent morphological stasis of Meioglossus may exemplify a potentially common evolutionary 'dead-end' scenario, where groups with highly miniaturized and simplified body plan lose their ability to diversify morphologically.

Coral Reef Water Microbial Communities of Jardines de la Reina, Cuba.

Globally, coral reef ecosystems are undergoing significant change related to climate change and anthropogenic activities. Yet, the Cuban archipelago of Jardines de la Reina (JR) has experienced fewer stressors due to its geographical remoteness and high level of conservation. This study examines the surface and benthic reef water microbial communities associated with 32 reef sites along the JR archipelago and explores the relationship between the community composition of reef microorganisms examined using bacterial and archaeal small subunit ribosomal RNA gene (16S rRNA gene) sequencing compared to geographic, conservation/protection level, environmental, physicochemical, and reef benthic and pelagic community features. Reef nutrient concentrations were low and microbial communities dominated by picocyanobacteria and SAR11 and SAR86 clade bacteria, characteristic of an oligotrophic system. Reef water microbial community alpha and beta diversity both varied throughout the archipelago and were strongly related to geography. Three sites in the western archipelago showed unique microbial communities, which may be related to the hydrogeography and influences of the channels linking the Ana Maria gulf with the Caribbean Sea. Overall, this work provides the first extensive description of the reef microbial ecology of the Caribbean's 'Crown Jewel' reef system and a framework to evaluate the influence of ongoing stressors on the reef microorganisms.

Historical changes of sediments and mollusk assemblages in the Gulf of Batabanó (Caribbean Sea) in the twentieth century.

The first paleoecological reconstruction of the biogeochemical conditions of the Gulf of Batabanó, Caribbean Sea was performed from (210)Pb-dated sediment cores. Depth profiles of 20 major elements and trace metals, organic compounds, grain size, and mollusk assemblage composition were determined from 9 stations encompassing unconsolidated sediments in the gulf. Spatial heterogeneity was evident for the geochemistry of sediments and for the mollusk assemblage composition. Our reconstruction indicates that pollution is not a critical threat to the ecosystem, although a slight historical increase of lead enrichment factor was detected probably due to long-range atmospheric fallout. Mollusk assemblages were composed by 168 species belonging to 59 families and no temporal trends in the species diversity or assemblage composition were detected, suggesting no depletion of diversity or habitat loss. Other signals of habitat loss such as changes in organic budget or increase of fine sediment fraction were absent or weak. Nitrogen retained in sediments changed by <1% in the century, indicating no historical events of eutrophication or oligotrophication in the gulf. Historical decrease of fine sediment fraction in the eastern sector would be linked to modifications in sedimentation rate, land use, and/or particle transport from the shelf border; this also suggests that both sectors have different sedimentary dynamics. Although, on theoretical grounds, historical fishery may have caused deleterious ecosystem effects by overexploitation of spiny lobster stocks, no evidence of habitat degradation or loss, caused by fisheries, could be detected.

Coral reef fish assemblages exhibit signs of depletion in two protected areas from the eastern of Los Canarreos archipelago (Cuba, Caribbean Sea).

Understanding the impact of marine protected areas on the distribution and composition of fishes is key to the protection and management of coral reef ecosystems, and especially for fish-based activities such as SCUBA diving and recreational fishing. The aim of this research is to compare the ichthyofauna structure in three areas in the eastern part of Los Canarreos archipelago in Cuba with different management schemes: Cayo Campos-Cayo Rosario Fauna Refuge (CCCR), Cayo Largo Ecological Reserve (CL) and non-protected area (nMPA), and considering habitat differences and depth variation. A total of 131 video transects were conducted using diver operated stereo-video (stereo-DOV) in November, 2015 in backreef and forereef along the CCCR, CL and the adjacent nMPA. We recorded 84 species and 27 functional groups suggesting high complementarity of functions. Several multispecies schools were observed along surveys, which explain the biomass peaks in some sites, mainly for Lutjanidae, Haemulidae and Carangidae. A concerning issue was the bare representation of critical functional groups and threatened species. The effect of sites nested within habitats was significant and the most important driver structuring fish assemblages, while MPA condition was not evident. Favorable habitat features (habitat heterogeneity and surrounding coastal ecosystems) are likely enhancing fish assemblages and counteracting the effects of pouching derived from insufficient management. We recommend immediate actions within a strategy of precautionary management including, but not limited to, the appointment of staff for the administration of CL, frequent monitoring and effective enforcement.

Mollusk death assemblages in 210Pb-dated marine sediment cores reveal recent biotic changes in the Gulf of Guanahacabibes, NW Cuba.

We investigated stratigraphic changes in mollusk death assemblages and geochemistry in sediment cores from four seagrass beds and one unvegetated site in the Gulf of Guanahacabibes (GG), NW Cuba. There was a transition from mangrove to seagrass beds, associated with sea level rise ∼6000 years ago. Sediment accumulation rates during the last century showed a general rise, but increased sharply after ∼1980, likely because of human activities. The GG displayed overall high mollusk γ-diversity, and our estimate of 189 species is biased toward the low end. High ß-diversity was driven by inter-site differences in grain size, vegetation cover, and nutrient input. Spatial heterogeneity within the basin influenced downcore abundance and diversity metrics, highlighting the influence of local drivers. Herbivorous gastropods dominated in seagrass beds and suspension feeder bivalves were dominant on sandy bottom. In the top parts of cores, species richness declined at two sites that were subject to high, human-mediated bulk sedimentation rates and eutrophication. Conservation measures are needed to preserve this hotspot of marine diversity.

Discovery and quantification of anaerobic nitrogen metabolisms among oxygenated tropical Cuban stony corals.

Coral reef health depends on an intricate relationship among the coral animal, photosynthetic algae, and a complex microbial community. The holobiont can impact the nutrient balance of their hosts amid an otherwise oligotrophic environment, including by cycling physiologically important nitrogen compounds. Here we use 15N-tracer experiments to produce the first simultaneous measurements of ammonium oxidation, nitrate reduction, and nitrous oxide (N2O) production among five iconic species of reef-building corals (Acropora palmata, Diploria labyrinthiformis, Orbicella faveolata, Porites astreoides, and Porites porites) in the highly protected Jardines de la Reina reefs of Cuba. Nitrate reduction is present in most species, but ammonium oxidation is low potentially due to photoinhibition and assimilatory competition. Coral-associated rates of N2O production indicate a widespread potential for denitrification, especially among D. labyrinthiformis, at rates of ~1 nmol cm-2 d-1. In contrast, A. palmata displays minimal active nitrogen metabolism. Enhanced rates of nitrate reduction and N2O production are observed coincident with dark net respiration periods. Genomes of bacterial cultures isolated from multiple coral species confirm that microorganisms with the ability to respire nitrate anaerobically to either dinitrogen gas or ammonium exist within the holobiont. This confirmation of anaerobic nitrogen metabolisms by coral-associated microorganisms sheds new light on coral and reef productivity.

A checklist of aquatic nematodes from Cuban Archipelago.

The diversity of free-living aquatic nematodes is largely unknown for the Gulf of Mexico and Caribbean Sea. The Cuban Archipelago is an important part of this because of its large area and diversity of habitats. We analyzed the free-living nematodes from 83 sites from seven aquatic habitats around Cuba, to produce a checklist for many habitats, including seagrass meadows, coral degradation zones, algal turf, bare sands, unvegetated muds, freshwater and anchihaline caves, and deep-sea sediments. The checklist contains 469 species, 229 genera, 50 families, and 9 orders. Chromadorida, Enoplida, and Monhysterida were the best represented orders with 112, 100, and 83 species respectively. The most abundant species were Euchromadora vulgaris, Terschellingia longicaudata, Desmodora pontica, Sabatieria pulchra , and Epsilonema sp. Most of the listed species were new records for the region. There were differences in the number of species recorded in each habitat type, with seagrass meadows having 280 species, coral degradation zones having 139 species, deep waters having 116 species, algal turf having 114 species, bare sands having 100 species, unvegetated muds having 78 species, freshwater caves having 19 species, anchihaline caves having 16 species, and freshwater streams having 6 species. The checklist is the most comprehensive recent report of the diversity of free-living nematodes in the regions of Gulf of Mexico and Caribbean Sea. The reported diversity is higher than many other regional checklists likely reflecting the intense sampling effort and the variety of microhabitats in Cuban Archipelago.

Diversity patterns of free-living nematode assemblages in seagrass beds from the Cuban archipelago (Caribbean Sea).

Diversity patterns of free-living marine nematodes in tropical seagrass beds are understudied. Here, we describe the species richness and assemblage composition of nematodes in 13 seagrass sites covering the whole Cuban archipelago. Nematodes were collected from Thalassia testudinum seagrass beds and identified to species level. We provide a checklist of nematode species from seagrass beds. The species richness of nematode assemblages is high, with 215 species, 138 genus, 35 families, seven orders and two classes. That γ-diversity is higher than other studies and points to seagrass beds as diversity hotspots of free-living marine nematodes. Local species richness in seagrass bed sites is about 57 ± 17 species and broadly similar across the sites, despite the environmental heterogeneity. The geographical distance plays a weak, but significant, role on the decay of similarity likely affected by limited dispersal of nematodes. The pairwise similarity values, related to poor-coloniser nematodes, were twice more affected by the distance than those related to good-colonisers, possibly due to differential success of transport and settlement.

Chronic PAH exposures and associated declines in fish health indices observed for ten grouper species in the Gulf of Mexico.

Ten grouper species grouper (n = 584) were collected throughout the Gulf of Mexico (GoM) from 2011 through 2017 to provide information on hepatobiliary polycyclic aromatic hydrocarbon (PAH) concentrations in the aftermath of the Deepwater Horizon (DWH) oil spill. Liver and bile samples were analyzed for PAHs and their metabolites using triple quadrupole mass spectrometry (GC/MS/MS) and high-performance liquid chromatography with fluorescence detection (HPLC-F), respectively. Data were compared among species and sub-regions of the GoM to understand spatiotemporal exposure dynamics in these economically and ecologically important species. Significant differences in the composition and concentrations of PAHs were detected spatially, over time and by species. The West Florida Shelf, Cuba coast and the Yucatan Shelf had a greater proportion of the pyrogenic PAHs in their livers than the other regions likely due to non-oil industry related sources (e.g., marine vessel traffic) in the regional composition profiles. Mean liver PAH concentrations were highest in the north central region of the GoM where DWH occurred. Biliary PAH concentrations and health indicator biometrics initially decrease during the first three years following the DWH oil spill but significantly increased thereafter. Increased exposures are likely explained by the resuspension of residual DWH oil as well as continued inputs from natural (e.g., seeps) sources and other anthropogenically derived sources (e.g., riverine runoff, other oil spills, and leaking oil and gas infrastructure). The increasing trend in PAH concentrations in the bile and liver of grouper species in the north central region of the GoM post-DWH suggest continued chronic exposures, however the critical stage at which permanent, irreparable damage may occur is unknown. Long-term monitoring of PAH levels and associated fish health biomarkers is necessary to evaluate impacts of chronic exposures, particularly in regions subject to intensive oil extraction activities.

A First Comprehensive Baseline of Hydrocarbon Pollution in Gulf of Mexico Fishes.

Despite over seven decades of production and hundreds of oil spills per year, there were no comprehensive baselines for petroleum contamination in the Gulf of Mexico (GoM) prior to this study. Subsequent to the 2010 Deepwater Horizon (DWH) spill, we implemented Gulf-wide fish surveys extending over seven years (2011-2018). A total of 2,503 fishes, comprised of 91 species, were sampled from 359 locations and evaluated for biliary polycyclic aromatic hydrocarbon (PAH) concentrations. The northern GoM had significantly higher total biliary PAH concentrations than the West Florida Shelf, and coastal regions off Mexico and Cuba. The highest concentrations of biliary PAH metabolites occurred in Yellowfin Tuna (Thunnus albacares), Golden Tilefish (Lopholatilus chamaeleonticeps), and Red Drum (Sciaenops ocellatus). Conversely, biliary PAH concentrations were relatively low for most other species including economically important snappers and groupers. While oil contamination in most demersal species in the north central GoM declined in the first few years following DWH, more recent increases in exposure to PAHs in some species suggest a complex interaction between multiple input sources and possible re-suspension or bioturbation of oil-contaminated sediments. This study provides the most comprehensive baselines of PAH exposure in fishes ever conducted for a large marine ecosystem.

Metabarcoding free-living marine nematodes using curated 18S and CO1 reference sequence databases for species-level taxonomic assignments.

High-throughput sequencing has the potential to describe biological communities with high efficiency yet comprehensive assessment of diversity with species-level resolution remains one of the most challenging aspects of metabarcoding studies. We investigated the utility of curated ribosomal and mitochondrial nematode reference sequence databases for determining phylum-specific species-level clustering thresholds. We compiled 438 ribosomal and 290 mitochondrial sequences which identified 99% and 94% as the species delineation clustering threshold, respectively. These thresholds were evaluated in HTS data from mock communities containing 39 nematode species as well as environmental samples from Vietnam. We compared the taxonomic description of the mocks generated by two read-merging and two clustering algorithms and the cluster-free Dada2 pipeline. Taxonomic assignment with the RDP classifier was assessed under different training sets. Our results showed that 36/39 mock nematode species were identified across the molecular markers (18S: 32, JB2: 19, JB3: 21) in UClust_ref OTUs at their respective clustering thresholds, outperforming UParse_denovo and the commonly used 97% similarity. Dada2 generated the most realistic number of ASVs (18S: 83, JB2: 75, JB3: 82), collectively identifying 30/39 mock species. The ribosomal marker outperformed the mitochondrial markers in terms of species and genus-level detections for both OTUs and ASVs. The number of taxonomic assignments of OTUs/ASVs was highest when the smallest reference database containing only nematode sequences was used and when sequences were truncated to the respective amplicon length. Overall, OTUs generated more species-level detections, which were, however, associated with higher error rates compared to ASVs. Genus-level assignments using ASVs exhibited higher accuracy and lower error rates compared to species-level assignments, suggesting that this is the most reliable pipeline for rapid assessment of alpha diversity from environmental samples.

Spatial and temporal variations of meiofaunal communities from the western sector of the Gulf of Batabanó, Cuba: II. Seagrass systems.

The meiofauna from seagrass meadows in the western sector of the Gulf of Batabanó, Cuba were studied to describe the spatial and temporal variations in community structure. Replicated cores were taken in three locations (arranged in m- and km-scales) and in two seasons (dry and wet). The meiofauna (metazoans between 500 and 45 microm) were identified to major taxa. Temporal changes in the meiofaunal communities could not be detected and they are not linked to the subtle seasonal changes in the water column. A larger variation in community structure was observed in the spatial m-scale (among cores in a station) probably accredited to heterogeneity of microenvironment and biological processes. A second source of variation in the km-scale (among locations) was identified relating to physical processes affecting seagrass meadows: marine currents and anthropogenic disturbances. Distribution patterns of meiofauna across locations coincide with one study from 20 years ago in seagrass beds (i.e. higher densities in area closer to break-shelf and diminution of fauna at southern of Pinar del Rio); however, cumulative anthropogenic disturbances on seagrass meadows would most likely explain the depletion of communities observed in our survey in comparison with decades ago. Estimates of meiofaunal density and richness of major taxa from our study (and other areas from the Cuban shelf) are consistently lower than other temperate and tropical sites; possibly caused by low primary productivity due to narrow tidal amplitude and oligotrophic waters.

Phylogeny and biogeography of the scaleless scale worm Pisione (Sigalionidae, Annelida).

Pisione is a scaleless group of small scale worms inhabiting sandy bottoms in shallow marine waters. This group was once considered rare, but now 45 described species can be characterized, among others, by their paired, segmental copulatory organs (one to multiple external pairs), which display a complexity of various accessory structures. The evolutionary significance of these unique organs was suggested in the late 1960s, but has been heavily debated since the late 1990s and remains controversial. In the present paper, we study the internal relationships within Pisione, employing combined phylogenetic analyses of both molecular and morphological data from 16 terminals of Pisione, as well as two terminals of Pisionidens, and eight additional scale worms as outgroups. Our taxon sampling covers all geographical areas where the genus has been reported, as well as most of their morphological and copulatory variability, including representatives of the "africana," "remota," "crassa," and "papuensis" groups, established previously by Yamanishi. We hereby provide a first insight into the relationships of the genus, testing previously proposed hypotheses on the evolutionary significance of male copulatory structures within Pisione, while attempting to understand patterns of distribution. The phylogenetic analyses using maximum likelihood and Bayesian methods consistently recovered two large clades spanning the East Atlantic (including the Mediterranean) and the Indo-Pacific-West Atlantic, respectively. Character optimization on our trees revealed a high degree of homoplasy in both non-reproductive and sexual characters of Pisione, with buccal acicula found to be the sole apomorphy among the morphological features assessed herein, with none defining the biogeographical subclades within. Overall, our comparative analyses highlight the high degree of morphological variation in this widely distributed genus, rejecting previous assertions of an increasing number and complexity of copulatory structures across the genus.

One-century decline of mollusk diversity as consequence of accumulative anthropogenic disturbance in a tropical estuary (Cuban Archipelago).

In order to infer changes in sediments and mollusk assemblages for the last century, we used biogeochemical data from two 210Pb dated cores collected in Sagua La Grande estuary, Cuban Archipelago. We found evidences of cumulative anthropogenic disturbance during the last century, causing considerable depletion of mollusk assemblage diversity and enhancement of the dominance of deposit feeding species. The sequence of impacts assessed was i) eutrophication due to nutrient releases from urban settlements, ii) habitat alteration due to water channeling and damming, and iii) mercury pollution. These successive impacts caused a steady diversity depletion from ca. 70 mollusk species in 1900 to less than five in 2010. Only two species persisted in the estuary: Nuculana acuta and Finella dubia. Hurricanes did not impact the molluscan fauna in the long term. The effects of the anthropogenic impacts suggest that the resilience of this estuarine system is compromised.

Two-year temporal response of benthic macrofauna and sediments to hypoxia in a tropical semi-enclosed bay (Cienfuegos, Cuba) / Respuesta temporal durante dos años de la macrofauna bentónica y sedimentos a la hipoxia en una bahía semi-cerrada tropical (Cienfuegos, Cuba)

AbstractHypoxia is the depletion of dissolved oxygen below 2 mg O2/L. Relatively few studies on hypoxia and its effects on benthic macrofauna have been done in tropical marine ecosystems. This study describes the temporal response of the water column, sediments and macrofauna to seasonal hypoxia in a semi-enclosed bay (Cienfuegos, Caribbean Sea). The Calisito site was sampled monthly from June 2010 until February 2012, yielding 21 sampling times. At each sampling event water and sediment samples were collected for measuring the abiotic variables (temperature, salinity, dissolved oxygen, nutrients, redox potential discontinuity, silt/clay and organic matter content) and macrofauna (abundance and species richness). Temperature and surface salinity followed a typical temporal pattern during the summer/rainy and the winter/dry periods. Salinity stratification occurred in the rainy period, lasting three months in 2010 and six months in 2011. The bottom water dissolved oxygen indicated hypoxic and anoxic events during the wet periods of 2010 and 2011 associated with salinity stratification, low hydrodynamics and oxidation of the accumulated organic matter. Over the study period, 817 individuals were collected and identified. Polychaetes were the dominant group in terms of abundance (57 % of total) followed by mollusks (41%). Hypoxia (and occasionally anoxia) caused strong deleterious effects on the abundance and species richness of macrofaunal communities in the study site. The most abundant polychaetes were opportunistic species with high tolerance to hypoxic conditions: Prionospio steenstrupi, Polydora sp.and Paraprionospio pinnata. Most of them colonized relatively fast once hypoxia ended. Persistent species such as Caecum pulchellum and Parvanachis obesa were present during hypoxia with fluctuating densities and apparently recover to higher abundances when normoxic conditions are re-established. Macoma tenta and Tellina consobrina colonized approximately 1-2 months later than the first polychaete peak during normoxia. Probably, the deleterious effects of hypoxia on the macrofauna were intensified by negative interspecific relationships such as competition by suitable space and predation. The recolonization of macrofauna depended possibly on local transport by currents within the bay because the connection with the Caribbean Sea is relatively limited. In summary, seasonal hypoxia in Cienfuegos Bay influences the water and sediment geochemistry and reduces both the abundance and diversity of macrofauna.

ResumenLa hipoxia es la disminución del oxígeno disuelto por debajo de 2 mg O2/L. Relativamente pocos estudios sobre la hipoxia y sus efectos en la macrofauna bentónica han sido realizados en ecosistemas marinos tropicales. Este estudio describe la respuesta temporal de la columna de agua, los sedimentos y la macrofauna a la hipoxia estacional en una bahía semicerrada en Cienfuegos, Mar Caribe. La estación de Calisito fue muestreada mensualmente de Junio 2010 a Febrero 2012, resultando en 21 eventos de muestreo. En cada oportunidad se recolectaron muestras de agua y sedimento para la medición de las variables abióticas (temperatura, salinidad, oxígeno disuelto, discontinuidad del potencial redox, contenido de limo/arcilla y materia orgánica) y macrofauna (abundancia y riqueza de especies). La temperatura y la salinidad superficial presentaron un patrón temporal típico durante los periodos de verano/ lluvia e invierno/seca. Ocurrió estratificación por salinidad en el periodo lluvioso, que duró tres meses en 2010 y seis en 2011. El oxígeno disuelto de fondo indicó eventos de hipoxia y anoxia durante los periodos lluviosos de 2010 y 2011 asociados con la estratificación de la columna de agua, la hidrodinámica lenta y la oxidación de la materia orgánica acumulada. Esto causó efectos deletéreos en la abundancia y diversidad de la macrofauna. Los poliquetos más abundantes fueron especies oportunistas con alta tolerancia a condiciones hipóxicas: Prionospio steenstrupi, Polydora sp. y Paraprionospio pinnata. La mayoría de estos colonizaron relativamente rápido una vez que terminó la hipoxia. Especies persistentes como Caecum pulchellum y Parvanachis obesa estuvieron presentes durante la hipoxia con densidades fluctuantes y aparentemente recuperaron sus abundancias cuando las condiciones normóxicas fueron re-establecidas. Macoma tenta y Tellina consobrina colonizaron 1-2 meses después que el primer pico de poliquetos durante la normoxia.Probablemente, los efectos deletéreos de la hipoxia en la macrofauna fueron intensificados por relaciones interespecíficas negativas como la competencia por espacio y la depredación. La recolonización de la macrofauna dependió probablemente del transporte local por las corrientes dentro de la bahía pues la conexión con el mar Caribe es relativamente limitada. En resumen, la hipoxia estacional en la bahía de Cienfuegos influencia la geoquímica del agua y los sedimentos y reduce la abundancia y diversidad de la macrofauna.