Niche divergence by deep‐sea octocorals in the genus Callogorgia across the continental slope of the Gulf of Mexico

Environmental variables that are correlated with depth have been suggested to be among the major forces underlying speciation in the deep sea. This study incorporated phylogenetics and ecological niche models (ENM) to examine whether congeneric species of Callogorgia (Octocorallia: Primnoidae) occupy different ecological niches across the continental slope of the Gulf of Mexico (GoM) and whether this niche divergence could be important in the evolution of these closely related species. Callogorgia americana americana, Callogorgia americana delta and Callogorgia gracilis were documented at 13 sites in the GoM (250–1000 m) from specimen collections and extensive video observations. On a first order, these species were separated by depth, with C. gracilis occurring at the shallowest sites, C. a. americana at mid‐depths and C. a. delta at the deepest sites. Callogorgia a. delta was associated with areas of increased seep activity, whereas C. gracilis and C. a. americana were associated with narrow, yet warmer, temperature ranges and did not occur near cold seeps. ENM background and identity tests revealed little to no overlap in ecological niches between species. Temporal calibration of the phylogeny revealed the formation of the Isthmus of Panama was a vicariance event that may explain some of the patterns of speciation within this genus. These results elucidate the potential mechanisms for speciation in the deep sea, emphasizing both bathymetric speciation and vicariance events in the evolution of a genus across multiple regions.     

Length–weight relationships of 11 deep‐sea fishes from the western Bay of Bengal and Andaman waters,India

Length–weight relationships (LWRs) are presented for 11 deep‐sea fishes caught in the western Bay of Bengal and Andaman waters during August 2010 using a 38 m high speed demersal trawl II (HSDT II, crustacean version, codend mesh size 40 mm) and a 45.6 m Expo model demersal trawl (codend mesh size 30 mm). The b values ranged from 2.34 to 3.3 and the coefficient of variation (r²) ranged from .82 to .98. LWR estimates of eight deep‐sea fishes are provided for the first time. The estimated LWR values were compared with the Bayesian LWR estimates available in FishBase, based on models developed to improve the accuracy and predictability of species‐specific growth parameters of data‐poor species.     

Molecular Phylogeny and Surface Morphology of Thiriotia hyperdolphinae n. sp. and Cephaloidophora oradareae n. sp. (Gregarinasina,Apicomplexa) Isolated from a Deep Sea Oradarea sp. (Amphipoda) in the West Pacific

In an effort to broaden our understanding of the biodiversity and distribution of gregarines infecting crustaceans, this study describes two new species of gregarines, Thiriotia hyperdolphinae n. sp. and Cephaloidophora oradareae n. sp., parasitizing a deep sea amphipod (Oradarea sp.). Amphipods were collected using the ROV Hyper‐Dolphin at a depth of 855 m while on a cruise in Sagami Bay, Japan. Gregarine trophozoites and gamonts were isolated from the gut of the amphipod and studied with light and scanning electron microscopy, and phylogenetic analysis of 18S rDNA. Thiriotia hyperdolphinae n. sp. was distinguished from existing species based on morphology, phylogenetic position, as well as host niche and geographic locality. Cephaloidophora oradareae n. sp. distinguished itself from existing Cephaloidophora, based on a difference in host (Oradarea sp.), geographic location, and to a certain extent morphology. We established this latter new species with the understanding that a more comprehensive examination of diversity at the molecular level is necessary within Cephaloidophora. Results from the 18S rDNA molecular phylogeny showed that T. hyperdolphinae n. sp. was positioned within a clade consisting of Thiriotia spp., while C. oradareae n. sp. grouped within the Cephaloidophoridae. Still, supplemental genetic information from gregarines infecting crustaceans will be needed to better understand relationships within this group of apicomplexans.     

Thaliacean distribution and abundance in the northern part of the Levantine Sea (Crete and Cyprus) during the eastern Mediterranean climatic transient, and a comparison with the western Mediterranean basin

First results are presented on the composition, abundance and vertical distribution of the thaliacean fauna in the Levantine basin obtained from stratified tows at three deep-sea sites in 1993: SE off Crete, and SW and NE off Cyprus. Samples with a 10 m²-MOCNESS (mesh size 1.67 mm) were poor in species and specimens as compared to samples with a 1 m²-double-MOCNESS (0.333 mm). Of the 12 species identified, six species belonged to the most abundant Doliolida, predominated by the phorozooids of Doliolum nationalis, five species belonged to the Salpida and one to the Pyrosomatida. Thaliaceans, most abundant by species and numbers SE off Crete, comprised ≤0.2% of the local mesozooplankton standing stocks. Presumably, they did not contribute substantially to the vertical flux generating the locally increased biomass and activity of the microbial benthos at the 4,300 m deep Cretean site. Most doliolids and salps were collected from the epipelagic and upper mesopelagic layers, and appeared to be most abundant close to the deep maximum of chlorophyll-a. Samples below 150 m were rarely rich in specimens, although two species performed diel migrations from the mesopelagic zone into the surface layers where some vertical segregation was evidenced. The aspect of niche separation is discussed.     

Distribution of <Emphasis Type="Italic">Calanus</Emphasis> populations in a glaciated fjord in the Arctic (Hornsund,Spitsbergen)—the interplay between biological and physical factors

The distribution and diversity of copepods of the genus Calanus were investigated in Hornsund Fjord (on the southwest coast of Spitsbergen) in summer 2001. The Bhattacharya method was used to sort individuals by species based on their prosome length. The established prosome length boundary values for the Calanus copepodid stages coincided with those defined for the Calanus species from Kongsfjorden (on the northwest coast of Spitsbergen). The predominant species in the main and inner fjord basins was Calanus glacialis, whereas Calanus finmarchicus was the prevailing species outside Hornsund. Younger copepodid stages (CI–CIII) of both species concentrated in the surface water layers (0–50∼70 m), while older copepodids (CIV–CVI females) that were ready for wintering stayed in deep layers (50∼70 m to bottom). Calanus hyperboreus was present in low numbers, predominantly as CIV, and in Hornsund deep water layers. The distribution and diversity of Calanus species complied with the notion that the marine fauna in Hornsund is of a more Arctic character than in Kongsfjorden, a fjord 260 km to the north on the west coast of Spitsbergen.     

Free‐living calamyzin chrysopetalids (Annelida) from methane seeps,anoxic basins,and whale falls

Members of Calamyzinae, a clade of free‐living and ectoparasitic chrysopetalids, are mainly associated with deep‐sea chemosynthetic environments. The three currently known free‐living calamyzin species are placed in Vigtorniella. A new free‐living calamyzin species similar to these is described here. Phylogenetic analyses of Calamyzinae using mitochondrial (cytochrome c oxidase subunit I and 16S rDNA) and nuclear (Histone H3 and 18S rDNA) loci showed that Vigtorniella and the new species form a grade with respect to an ectoparasitic clade, requiring two new genera to be erected. All free‐living calamyzins show a similar anterior end and chaetal morphology. Micospina auribohnorum gen. et sp. nov. is described for the small‐bodied new species from deep‐sea whale falls off California and methane seeps off Costa Rica. The maximum‐likelihood and Bayesian analyses show Micospina gen. nov. as sister to the ectoparasitic clade. Boudemos gen. nov. is named for the clade of two larger‐bodied species: Boudemos flokati gen. et comb. nov. and Boudemos ardabilia gen. et comb. nov., which is sister group to all other Calamyzinae. Vigtorniella is retained for the type species, Vigtorniella zaikai (Kiseleva, 1992), with the adults found amongst bacterial mats at the boundary of the hydrogen sulphide zone in the Black Sea. Micospina gen. nov., Boudemos gen. nov., and Vigtorniella form a grade of free‐living taxa that is associated with feeding on organic‐enriched sediments, and the latter two taxa display ontogenetic jaw change. Jaws are absent in Micospina auribohnorum gen. et sp. nov. and most of the calamyzin clade of parasitic forms.     

Length–weight relationship of six deep‐sea fish species from the shelf regions of western Bay of Bengal and Andaman waters

Length–weight relationships for six deep‐sea fish species inhabiting the shelf regions of the western Bay of Bengal and the waters of Andaman and Nicobar of India are presented. Samples were collected using high‐speed demersal trawl and expo demersal trawl nets at depths of 231–514 m in August and September 2010. The b values in the analyses ranged from 3.05 to 3.31, showing a mean and median value of 3.21 (SE ± 0.039, SD ± 0.097) and 3.2, respectively. Comparisons of b values with earlier estimations confirm the presence of spatial and temporal variations in the length–weight relations among the species. Coefficient of determination scores ranged from 0.94 to 0.97, indicating robustness of the samples analysed. This study provides the first estimates of length–weight relationships for four of the deep‐sea fishes, enriching the understanding of the growth patterns and population dynamics of these less‐studied deep‐sea fishery resources in Indian Ocean waters.     

Visual perception of light organ patterns in deep‐sea shrimps and implications for conspecific recognition

Darkness and low biomass make it challenging for animals to find and identify one another in the deep sea. While spatiotemporal variation in bioluminescence is thought to underlie mate recognition for some species, its role in conspecific recognition remains unclear. The deep‐sea shrimp genus, Sergestes sensu lato (s.l.), is one group that is characterized by species‐specific variation in light organ arrangement, providing us the opportunity to test whether organ variation permits recognition to the species level. To test this, we analyzed the visual capabilities of three species of Sergestes s.l. in order to (a) test for sexual dimorphism in eye‐to‐body size scaling relationships, (b) model the visual ranges (i.e., sighting distances) over which these shrimps can detect intraspecific bioluminescence, and (c) assess the maximum possible spatial resolution of the eyes of these shrimps to estimate their capacity to distinguish the light organs of each species. Our results showed that relative eye size scaled negatively with body length across species and without sexual dimorphism. Though the three species appear capable of detecting one another's bioluminescence over distances ranging from < 1 to ~6 m, their limited spatial resolution suggests they cannot resolve light organ variation for the purpose of conspecific recognition. Our findings point to factors other than conspecific recognition (e.g., neutral drift, phenotypic constraint) that have led to the extensive diversification of light organs in Sergestes s.l and impart caution about interpreting ecological significance of visual characters based on the resolution of human vision. This work provides new insight into deep‐sea animal interaction, supporting the idea that—at least for these mesopelagic shrimps—nonvisual signals may be required for conspecific recognition.     

Surface-associated diatoms from marine habitats at Cape Evans,Antarctica, including the first record of living <Emphasis Type="Italic">Eunotogramma marginopunctatum</Emphasis>

The diversity of diatom communities found epiphytically on red macroalgae collected under the sea ice at Cape Evans, Antarctica, was studied microscopically. Comparison of communities along a depth profile showed that species diversity decreased as depth below sea ice increased. Dominant taxa also changed with depth, with Cocconeis fasciolata dominant at 10 and 15 m, Porosira glacialis at 20 m and Eunotogramma marginopunctatum at 25 m. Epiphytic communities were also compared to sympagic and sediment-associated communities collected from a single depth. Species richness was greatest for communities associated with surface sediments while the poorest was found in sympagic, or sea-ice, communities. E. marginopunctatum, previously only described from fossil material, was found associated epiphytically on macroalgae from 20 to 25 m below sea ice.     

Mantoniella beaufortii and Mantoniella baffinensis sp. nov. (Mamiellales,Mamiellophyceae), two new green algal species from the high arctic1

Members of the class Mamiellophyceae comprise species that can dominate picophytoplankton diversity in polar waters. Yet, polar species are often morphologically indistinguishable from temperate species, although clearly separated by molecular features. Here we examine four Mamiellophyceae strains from the Canadian Arctic. The 18S rRNA and Internal Transcribed Spacer 2 (ITS2) gene phylogeny place these strains within the family Mamiellaceae (Mamiellales, Mamiellophyceae) in two separate clades of the genus Mantoniella. ITS2 synapomorphies support their placement as two new species, Mantoniella beaufortii and Mantoniella baffinensis. Both species have round green cells with diameter between 3 and 5 μm, one long flagellum and a short flagellum (~1 μm) and are covered by spiderweb-like scales, making both species similar to other Mantoniella species. Morphologically, M. beaufortii and M. baffinensis are most similar to the cosmopolitan M. squamata with only minor differences in scale structure distinguishing them. Screening of global marine metabarcoding data sets indicates M. beaufortii has only been recorded in seawater and sea ice samples from the Arctic, while no environmental barcode matches M. baffinensis. Like other Mamiellophyceae genera that have distinct polar and temperate species, the polar distribution of these new species suggests they are cold or ice-adapted Mantoniella species.     

Ultrastructure and Systematics of Two New Species of Dinoflagellate,Paragymnodinium Asymmetricum sp. nov. and Paragymnodinium Inerme sp. nov. (Gymnodiniales,Dinophyceae)1

The genus Paragymnodinium currently includes two species, P. shiwhaense and P. stigmaticum, that are characterized by mixotrophic nutrition and the possession of nematocysts. In this study, two new dinoflagellates belonging to this genus were described based on observations using LM, SEM, and TEM together with a molecular analysis. Cells of P. asymmetricum sp. nov., isolated from Nha Trang Beach, Vietnam, were 7.9–12.6 μm long and 4.7–9.0 μm wide. The species showed no evidence of feeding behavior and was able to sustain itself phototrophically. Paragymnodinium asymmetricum shared many features with P. shiwhaense, including presence of nematocysts, absence of an eyespot, and a planktonic lifestyle, but was clearly distinguished by the asymmetric shape of the hyposome, possession of a single chloroplast, and its nutritional mode. Cells of P. inerme sp. nov., isolated from Jogashima, Kanagawa Pref, Japan, were 15.3–23.7 μm long and 10.9–19.6 μm wide. This species also showed no evidence of feeding behavior. Paragymnodinium inerme was similar to cells of P. shiwhaense in shape and planktonic lifestyle, but its nutritional mode was different. The presence of incomplete nematocysts was also a unique feature. A phylogenetic analysis inferred from concatenated SSU and LSU rDNA sequences recovered the two dinoflagellates in a robust clade with Paragymnodinium spp., within the clade of Gymnodinium sensu stricto. This evidence, together with their morphological similarities, made it reasonable to conclude that these two dinoflagellates are new species of Paragymnodinium.     

The mitochondrial genome of a sea anemone Bolocera sp. exhibits novel genetic structures potentially involved in adaptation to the deep‐sea environment

The deep sea is one of the most extensive ecosystems on earth. Organisms living there survive in an extremely harsh environment, and their mitochondrial energy metabolism might be a result of evolution. As one of the most important organelles, mitochondria generate energy through energy metabolism and play an important role in almost all biological activities. In this study, the mitogenome of a deep‐sea sea anemone (Bolocera sp.) was sequenced and characterized. Like other metazoans, it contained 13 energy pathway protein‐coding genes and two ribosomal RNAs. However, it also exhibited some unique features: just two transfer RNA genes, two group I introns, two transposon‐like noncanonical open reading frames (ORFs), and a control region‐like (CR‐like) element. All of the mitochondrial genes were coded by the same strand (the H‐strand). The genetic order and orientation were identical to those of most sequenced actiniarians. Phylogenetic analyses showed that this species was closely related to Bolocera tuediae. Positive selection analysis showed that three residues (31 L and 42 N in ATP6, 570 S in ND5) of Bolocera sp. were positively selected sites. By comparing these features with those of shallow sea anemone species, we deduced that these novel gene features may influence the activity of mitochondrial genes. This study may provide some clues regarding the adaptation of Bolocera sp. to the deep‐sea environment.     

Invertebrate population genetics across Earth's largest habitat: The deep‐sea floor

Despite the deep sea being the largest habitat on Earth, there are just 77 population genetic studies of invertebrates (115 species) inhabiting non‐chemosynthetic ecosystems on the deep‐sea floor (below 200 m depth). We review and synthesize the results of these papers. Studies reveal levels of genetic diversity comparable to shallow‐water species. Generally, populations at similar depths were well connected over 100s–1,000s km, but studies that sampled across depth ranges reveal population structure at much smaller scales (100s–1,000s m) consistent with isolation by adaptation across environmental gradients, or the existence of physical barriers to connectivity with depth. Few studies were ocean‐wide (under 4%), and 48% were Atlantic‐focused. There is strong emphasis on megafauna and commercial species with research into meiofauna, “ecosystem engineers” and other ecologically important species lacking. Only nine papers account for ~50% of the planet's surface (depths below 3,500 m). Just two species were studied below 5,000 m, a quarter of Earth's seafloor. Most studies used single‐locus mitochondrial genes revealing a common pattern of non‐neutrality, consistent with demographic instability or selective sweeps; similar to deep‐sea hydrothermal vent fauna. The absence of a clear difference between vent and non‐vent could signify that demographic instability is common in the deep sea, or that selective sweeps render single‐locus mitochondrial studies demographically uninformative. The number of population genetics studies to date is miniscule in relation to the size of the deep sea. The paucity of studies constrains meta‐analyses where broad inferences about deep‐sea ecology could be made.     

Vertical distribution of Stomoxys spp. (Diptera: Muscidae) in a rainforest area of Gabon

The vertical distribution of Stomoxys spp. was studied in a rainforest area, Ipassa‐Makokou biosphere reserve located in the Ivindo National Park of Gabon. From April to June 2006, Vavoua traps were set out during 15 consecutive days per month at different heights above ground level corresponding to vertical layers of rainforest: 50 cm, 10, 20 and 30 m. Stomoxys calcitrans, S. transvittatus, S. omega, S. niger niger and S. niger bilineatus were more abundant at near ground level (50 cm), whereas abundance of S. xanthomelas was greatest in traps higher (20 and 30 m) in the canopy. Fly abundance was significantly different among vertical layers of the forest (H = 36.91; P < 0.001, ddl = 3), and among species to another (H = 41.11, P < 0.001). Vertical distribution of fly species corroborates feeding behaviour as the identification of blood meal origins showed heterogeneity of feeding hosts. High densities of flies were also observed at 10 m, and most S. inornatus were captured at that level. These results show that Stomoxyine flies in this rainforest are present in all vertical layers, from the ground level to the canopy. Their ubiquity, regarding both their habitats and their hosts, should be taken into account if a vector control strategy is planned in this touristic area.     

Molecular and Morphological Characterization of Anncaliia azovica sp. n. (Microsporidia) Infecting Niphargogammarus intermedius (Crustacea,Amphipoda) from the Azov Sea

Five out of one hundred adults of Niphargogammarus intermedius caught at the Azov sea shore were found to be infected with microsporidia. The infection was found in the subcuticular fat body and myocytes. Parasites developed in direct contact with host cells, displayed a disporoblastic sporogony and diplokaryotic arrangement of nuclei at all stages. Spores were oval, 4.6–5.8 × 2.6–3.0 μm. Exospore appendages, vesicular–tubular secretions, and the anisofilar polar filament indicated a similarity to Anncaliia species. Sporont surfaces displayed ridges of amorphous material. Meronts and sporonts formed protoplastic extensions, similar to A. vesicularum and A. meligheti. Mature spores possessed a bipartite polaroplast. The polar tube was arranged in one row of 13–18 coils including 0–3 distal coils of lesser diameter. Partial sequencing of SSU, ITS, and LSU regions of rRNA gene (GenBank accessions: KY288064 – KY288065 ) confirmed this new species to be congeneric with A. algerae (# AF069063 ) and A. meligheti (# AY894423 ). The SSU gene of this novel microsporidium shared 99.4% sequence similarity to A. algerae and 98.9% to A. meligheti. Genes for HSP70 and RPB1 amplified with primers designed for A. algerae orthologs displayed 99.7% and 97.4% similarity, respectively, between A. algerae and the novel microsporidium. A new species, Anncaliia azovica, is described based on morphological and molecular characterization.     

Oractis bursifera sp. nov., an Arctic deep-sea anemone with peculiar invaginations of its oral disc (Cnidaria: Actiniaria)

In 1993, a small, inconspicuous sea anemone, Oractis bursifera sp. nov., was collected during cruise ARK IX/4 of RV “Polarstern” from 3000 m depth in the central Arctic deep sea (Amundsen Basin). Histological sections revealed a unique morphological trait in the form of ten sac-like invaginations of the oral disc lined with an unusual, presumably glandular, epithelium. Another remarkable observation relates to its sexual status: testicular follicles are present in the eight macromesenteries, with one of these mesenteries also harbouring a single, large, apparently mature oocyte. Whether these peculiarities are related to each other and may thus be part of a reproductive pattern in extreme habitats such as the Arctic deep sea is discussed. The peculiar characters of the new species confirm the extraordinary position of the genus Oractis McMurrich, 1893, and warrant the establishment of the new family Oractiidae within the endomyarian sea anemones. Accepted: 6 March 2000     

First records of sea snakes (Elapidae: Hydrophiinae) diving to the mesopelagic zone (>200 m)

Viviparous sea snakes (Elapidae: Hydrophiinae) are fully marine reptiles distributed in the tropical and subtropical waters of the Indian and Pacific Oceans. Their known maximum diving depth ranges between 50 and 100 m and this is thought to limit their ecological ranges to shallow habitats. We report two observations, from industry‐owned remotely operated vehicles, of hydrophiine sea snakes swimming and foraging at depths of approximately 250 m in the Browse Basin on Australia's North West Shelf, in 2014 and 2017. These observations show that sea snakes are capable of diving to the dim‐lit, cold‐water mesopelagic zone, also known as the ‘twilight’ zone. These record‐setting dives raise new questions about the thermal tolerances, diving behaviour and ecological requirements of sea snakes. In addition to significantly extending previous diving records for sea snakes, these observations highlight the importance of university‐industry collaboration in surveying understudied deep‐sea habitats.     

A New Heterolobosean Amoeboflagellate,Tetramitus dokdoensis n. sp., Isolated from a Freshwater Pond on Dokdo Island in the East Sea,Korea

The genus Tetramitus is a representative amoeboflagellate group within the Heterolobosea, and currently contains over a dozen species. Here, a new heterolobosean amoeboflagellate was isolated from a freshwater pond on Dokdo Island, Korea. The amoebae have eruptive pseudopodia, no uroidal filament, and a nucleus with a central nucleolus. The length and width of the amoebae are 15.5–28.0 μm and 5.4–12.6 μm, respectively. The flagellates are conical, with 4 flagella of equal length (~10 μm). There is a discrete rostrum in the subapical region of the flagellate form. The cyst has thin endo‐ and ectocyst layers and no cyst pores. The amoeba shows slow movement at 37 °C, but does not move at 42 °C under a light microscope. Phylogenies of the 18S rRNA gene and the ITS1‐5.8S rRNA gene‐ITS2 sequence show that the strain belongs to a subclade of Tetramitus that includes Tetramitus rostratus, Tetramitus waccamawensis and Tetramitus entericus, amongst others. Nonetheless, the strain is distinct from other species in both molecular phylogenetic trees. Thus the strain isolated from the Dokdo Island is proposed as a novel species, Tetramitus dokdoensis n. sp.     

Habitat-based submersible assessment of macro-invertebrate and groundfish assemblages at Heceta Bank, Oregon, from 1988 to 1990

Exploitation of groundfish off the U.S. Pacific coast reached maximum levels during the 1990s, resulting in severe declines in at least nine species of groundfish. From 1988 to 1990, we used the 2-man submersible Delta to make 42 dives and run replicate visual belt transects at six stations ranging from 67-360 m in depth at Heceta Bank on the outer continental shelf of Oregon. We identified four major habitats and associated benthic macroinvertebrate and groundfish assemblages: (1) shallow rock ridges and large boulders (< 100 m deep) dominated by basket stars, juvenile rockfishes, yelloweye rockfish, and lingcod; (2) mid-depth small boulder-cobbles (100-150 m) dominated by crinoids, brittle stars, rosethorn, pygmy/Puget sound, and canary rockfishes; (3) deep cobble (150-200 m) dominated by crinoids, brittle stars and various small rockfish species, and (4) deep mud slope (> 200 m) dominated by fragile urchins, sea cucumbers, shortspine thornyhead, and flatfishes. Although substantial interannual variation in groundfish abundance among seafloor types was evident in the 12 most abundant and/or commercially important fish taxa sampled, high variance resulted in statistically significant differences among years only in juvenile rockfishes. These data provide a baseline for future comparisons exploring long-term change this continental-shelf ecosystem.