Dancing between the devil and deep blue sea: the stabilizing effect of enemy-free and victimless sinks

Theoretical and empirical studies have shown that enemy–victim interactions in spatially homogenous environments can exhibit diverging oscillations which result in the extinction of one or both species. For enemy–victim models with overlapping generations, we investigate the dynamical implications of spatial heterogeneity created by enemy-free sinks or victimless sinks. An enemy-free sink is a behavioral, physiological or ecological state that reduces or eliminates the victim's vulnerability to the enemy but cannot sustain the victim population. For victims that move in an ideal-free manner, we prove that the inclusion of an enemy-free sink shifts the population dynamics from diverging oscillations to stable oscillations. During these stable oscillations, the victim disperses in an oscillatory manner between the enemy-free sink and the enemy-occupied patch. Enemy-free sinks with lower mortality rates exhibit oscillations with smaller amplitudes and longer periods. A victimless sink, on the other hand, is a behavioral, physiological or ecological state in which the enemy has limited (or no) access to its victims. For enemies that move in an ideal-free manner, we prove that victimless sinks also stabilize diverging oscillations. Simulations suggest that suboptimal behavior due to information gathering or learning limitations amplify oscillations for systems with enemy-free sinks and dampen oscillations for systems with victimless sinks. These results illustrate that the coupling of a sink created by unstable enemy–victim interactions and a sink created by unsuitable environmental conditions can result in population persistence at the landscape level.     

The invasive green crab and Japanese shore crab: behavioral interactions with a native crab species,the blue crab

Blue crabs, Callinectes sapidus (Rathbun), are an ecologically and commercially important species along the East coast of North America. Over the past century and a half, blue crabs have been exposed to an expanding set of exotic species, a few of which are potential competitors. To test for interactions with invasive crabs, juvenile C. sapidus males were placed in competition experiments for a food item with two common non-indigenous crabs, the green crab Carcinus maenas (L.) and the Japanese shore crab, Hemigrapsus sanguineus (De Haan). Agonistic interactions were evaluated when they occurred. In addition, each species’ potential to resist predators was examined by testing carapace strength. Results showed that C. maenas was a superior competitor to both C. sapidus and H. sanguineus for obtaining food, while the latter two species were evenly matched against each other. Regarding agonism, C. sapidus, was the “loser” a disproportionate number of times. C. sapidus carapaces also had a significantly lower breaking strength. These experiments suggest that both as a competitor, and as potential prey, juvenile blue crabs have some disadvantages compared with these common sympatric exotic crab species, and in areas where these exotics are common, juvenile native blue crabs may be forced to expend more energy in conflict that could be spent foraging, and may be forced away from prime food items toward less optimum prey.     

A new genus and two new species of Choniostomatidae (Copepoda) parasitic on two deep sea isopods

The systematic position of the little-deformed parasitic copepod family Nicothoidae to the much deformed family Choniostomatidae has been disputed for many years. The new genus Diexanthema gen. nov., based on Diexanthema desistoma sp. nov. and Diexanthema bathydiaita sp. nov. discovered on two deep-sea asellote isopod species, clearly links the family Nicothoidae (via male Diexanthema) to the family Choniostomatidae (via female Diexanthema). Based upon the extreme sexual dimorphism of this new genus, it is suggested that the family Nicothoidae be dissolved and its members placed within the family Choniostomatidae. It is concluded that the differences seen between these two parasitic groups is probably due more to host adaptations than reflecting any phylogenetic trend that is evolutionarily significant. It is felt that the deep-sea environment may serve as a refugium for archaic copepod forms which may possibly offer additional clues and insights into the phylogeny of the parasitic copepoda.
This investigation suggests that the life history and parasite/host relationships of Diexanthema desistoma are quite complicated and well geared to the life history of its isopod host, Chelator insignis.     

Highly migratory species in the new law of the sea

Abstract

The 1982 Convention on the Law of the Sea provides for coastal state sovereign rights over all living resources within the exclusive economic zone, but in a separate article makes special provision for so‐called highly migratory species (HMS) that are specifically identified in an Annex to the treaty. This paper examines the basic treaty provisions concerning all fisheries in the EEZ, highlighting the position of HMS, and then devotes detailed consideration to the regulation of fishing for HMS in light of Article 64 and other articles of the treaty. Discussion centers upon the requirements of Article 64, including the duty of cooperation and its subject matter, the meaning and consequence of failure to cooperate, the manner of cooperation under Article 64, and the application of the treaty to enclaves of high seas. Attention is given to the application to HMS of specific obligations under Articles 61 and 62, authority over HMS fishing within archipelagic waters, and the use of fish aggregation devices. There is also consideration of the relationship between the 1982 Convention and customary law regarding HMS, with particular attention to the position vis‐a‐vis the Convention of the United States as a nonsignatory who does not recognize coastal state jurisdiction over tuna but claims sovereign rights over all other highly migratory species within its recently proclaimed exclusive economic zone.     

Four new species of Desmosomatidae Sars, 1897 (Crustacea: Isopoda) from the deep sea of the Angola Basin

From a total of 104 species and 1741 individuals of Isopoda, the material from DIVA-1 (DIVersity of the Atlantic benthos) contained 364 specimens belonging to Desmosomatidae Sars, 1897. They are represented by 10 genera and 27 species. Most species (93%) are new to science. Four species are described in this study (Eugerdella theodori sp. nov., Desmosoma renatae sp. nov., Torwolia tinbiene sp. nov. and Momedossa longipedis sp. nov.). A discussion of the genera Desmosoma Sars, 1864, Momedossa Hessler, 1970 and Torwolia Hessler, 1970 is presented.     

Alternative prey: a mechanism for elimination of bacterial species by protozoa

Antibiotic-resistant strains of Salmonella typhimurium and Klebsiella pneumoniae died readily after their addition to raw sewage, but they grew in sterilized sewage. The decline was not a result of abiotic stresses, and because the bacteria were able to survive in large numbers for at least 15 days in solutions containing no organic nutrients, it was not a result of competition. Toxin production, bacteriophages, and Bdellovibrio sp. did not cause the disappearance of the two bacterial species. A decline was also evident if the sewage was first passed through a 3-micron (pore size) filter or treated with cycloheximide or cycloheximide plus nystatin, but protozoa developed under these conditions. Little or no decline occurred if the sewage was filtered and treated with the eucaryotic inhibitors before the addition of S. typhimurium or K. pneumoniae, and protozoa were not detected. S. typhimurium increased in abundance if cycloheximide, streptomycin, and erythromycin or large amounts of glucose were added to sewage. Tetrahymena thermophilus did not significantly reduce the population of S. typhimurium in buffer when the density of the bacterium was about 10(4)/ml. However, when more than 10(8) Enterobacter agglomerans cells per ml were added to the buffer, T. thermophilus reduced the abundance of E. agglomerans and S. typhimurium to 10(6) and 10/ml, respectively. The density of S. typhimurium was further decreased by a second increment of E. agglomerans cells. The disappearance of S. typhimurium and K. pneumoniae from sewage thus is the result of predation by protozoa.(ABSTRACT TRUNCATED AT 250 WORDS)     

Colonization of the deep sea by fishes

Analysis of maximum depth of occurrence of 11 952 marine fish species shows a global decrease in species number (N) with depth (x; m): log₁₀N = ?0·000422x + 3·610000 (r² = 0·948). The rate of decrease is close to global estimates for change in pelagic and benthic biomass with depth (?0·000430), indicating that species richness of fishes may be limited by food energy availability in the deep sea. The slopes for the Classes Myxini (?0·000488) and Actinopterygii (?0·000413) follow this trend but Chondrichthyes decrease more rapidly (?0·000731) implying deficiency in ability to colonize the deep sea. Maximum depths attained are 2743, 4156 and 8370 m for Myxini, Chondrichthyes and Actinopterygii, respectively. Endemic species occur in abundance at 7–7800 m depth in hadal trenches but appear to be absent from the deepest parts of the oceans, >9000 m deep. There have been six global oceanic anoxic events (OAE) since the origin of the major fish taxa in the Devonian c. 400 million years ago (mya ). Colonization of the deep sea has taken place largely since the most recent OAE in the Cretaceous 94 mya when the Atlantic Ocean opened up. Patterns of global oceanic circulation oxygenating the deep ocean basins became established coinciding with a period of teleost diversification and appearance of the Acanthopterygii. Within the Actinopterygii, there is a trend for greater invasion of the deep sea by the lower taxa in accordance with the Andriashev paradigm. Here, 31 deep‐sea families of Actinopterygii were identified with mean maximum depth >1000 m and with >10 species. Those with most of their constituent species living shallower than 1000 m are proposed as invasive, with extinctions in the deep being continuously balanced by export of species from shallow seas. Specialized families with most species deeper than 1000 m are termed deep‐sea endemics in this study; these appear to persist in the deep by virtue of global distribution enabling recovery from regional extinctions. Deep‐sea invasive families such as Ophidiidae and Liparidae make the greatest contribution to fish fauna at depths >6000 m.     

Evidence for Permo-Triassic colonization of the deep sea by isopods

The deep sea is one of the largest ecosystems on Earth and is home to a highly diverse fauna, with polychaetes, molluscs and peracarid crustaceans as dominant groups. A number of studies have proposed that this fauna did not survive the anoxic events that occurred during the Mesozoic Era. Accordingly, the modern fauna is thought to be relatively young, perhaps having colonized the deep sea after the Eocene/Oligocene boundary. To test this hypothesis, we performed phylogenetic analyses of nuclear ribosomal 18S and 28S and mitochondrial cytochrome oxidase I and 16S sequences from isopod crustaceans. Using a molecular clock calibrated with multiple isopod fossils, we estimated the timing of deep-sea colonization events by isopods. Our results show that some groups have an ancient origin in the deep sea, with the earliest estimated dates spanning 232–314 Myr ago. Therefore, anoxic events at the Permian–Triassic boundary and during the Mesozoic did not cause the extinction of all the deep-sea fauna; some species may have gone extinct while others survived and proliferated. The monophyly of the ‘munnopsid radiation’ within the isopods suggests that the ancestors of this group evolved in the deep sea and did not move to shallow-water refugia during anoxic events.     

Speedy speciation in a bacterial microcosm: new species can arise as frequently as adaptations within a species

Microbiologists are challenged to explain the origins of enormous numbers of bacterial species worldwide. Contributing to this extreme diversity may be a simpler process of speciation in bacteria than in animals and plants, requiring neither sexual nor geographical isolation between nascent species. Here, we propose and test a novel hypothesis for the extreme diversity of bacterial species—that splitting of one population into multiple ecologically distinct populations (cladogenesis) may be as frequent as adaptive improvements within a single population''s lineage (anagenesis). We employed a set of experimental microcosms to address the relative rates of adaptive cladogenesis and anagenesis among the descendants of a Bacillus subtilis clone, in the absence of competing species. Analysis of the evolutionary trajectories of genetic markers indicated that in at least 7 of 10 replicate microcosm communities, the original population founded one or more new, ecologically distinct populations (ecotypes) before a single anagenetic event occurred within the original population. We were able to support this inference by identifying putative ecotypes formed in these communities through differences in genetic marker association, colony morphology and microhabitat association; we then confirmed the ecological distinctness of these putative ecotypes in competition experiments. Adaptive mutations leading to new ecotypes appeared to be about as common as those improving fitness within an existing ecotype. These results suggest near parity of anagenesis and cladogenesis rates in natural populations that are depauperate of bacterial diversity.     

Mid-domain models as predictors of species diversity patterns: bathymetric diversity gradients in the deep sea

Geometric constraints represent a class of null models that describe how species diversity may vary between hard boundaries that limit geographic distributions. Recent studies have suggested that a number of large scale biogeographic patterns of diversity (e.g. latitude, altitude, depth) may reflect boundary constraints. However, few studies have rigorously tested the degree to which mid-domain null predictions match empirical patterns or how sensitive the null models are to various assumptions. We explore how variation in the assumptions of these models alter null depth ranges and consequently bathymetric variation in diversity, and test the extent to which bathymetric patterns of species diversity in deep sea gastropods, bivalves, and polychaetes match null predictions based on geometric constraints.
Range–size distributions and geographic patterns of diversity produced by these null models are sensitive to the relative position of the hard boundaries, the specific algorithms used to generate range sizes, and whether species are continuously or patchily distributed between range end points. How well empirical patterns support null expectations is highly dependent on these assumptions. Bathymetric patterns of species diversity for gastropods, bivalves and polychaetes differ substantially from null expectations suggesting that geometric constraints do not account for diversity–depth patterns in the deep sea benthos.     

Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores,including one new genus and three new species

The Hippasterinae is a subfamily within the Goniasteridae, consisting of five genera and 26 species, which occur in cold‐water settings ranging from subtidal to abyssal depths. All known genera were included in a cladistic analysis resulting in two most parsimonious trees, supporting the Hippasterinae as monophyletic. Our review supports Sthenaster emmae gen. et sp. nov. as a new genus and species from the tropical Atlantic and two new Evoplosoma species, Evoplosoma claguei sp. nov. and Evoplosoma voratus sp. nov. from seamounts in the North Pacific. Hippasteria caribaea is reassigned to the genus Gilbertaster, which previously contained a single Pacific species. Our analysis supports Evoplosoma as a derived deep water lineage relative to its continental‐shelf, shallow water sister taxa. The genus Hippasteria contains approximately 15 widely distributed, but similar‐looking species, which occur in the northern and southern hemispheres. Except for Gilbertaster, at least one species in each genus has been observed or is inferred to prey on deep‐sea corals, suggesting that this lineage is important to the conservation of deep‐sea coral habitats. The Hippasterinae shares several morphological similarities with Circeaster and Calliaster, suggesting that they may be related. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 266–301.     

深层海洋浮游植物研究综述

对深层海洋(200 m)浮游生物的研究是我国海洋浮游生物研究的前沿,已有的深层海洋浮游生物知识是我国相关研究的基础。浮游植物需要阳光进行光合作用,大多数浮游植物沉降到深海中会死亡,但有些浮游植物却能在深海中生存。综述深层海洋中存活的浮游植物的研究进展,为我国的研究提供参考。在深层海洋发现的浮游植物(2"m)种类不多,共有18种,最深分布在4000 m,颗石藻的丰度大于硅藻和甲藻,最大丰度为2220个/L,出现在500 m。在1911—1985年期间,曾经发现橄榄绿细胞,由于研究手段有限,一直没有对其进行分类学研究。1985年以后的研究怀疑这些橄榄绿细胞就是聚球藻蓝细菌,聚球藻蓝细菌在太平洋、大西洋和地中海的几个站位能分布到2750 m,在深层海洋的最大丰度为3.5×105个/mL,出现在800 m。在许多海区200—700 m发现有浮游植物色素荧光的高值,但对引起高值的原因尚不清楚。浮游植物生物量在深层海洋的垂直剖面研究不多,已有的研究表明浮游植物生物量在300 m以深为0.001—0.1"g C/L,且其随深度增加而降低。     

Alternative prey: a mechanism for elimination of bacterial species by protozoa.

Antibiotic-resistant strains of Salmonella typhimurium and Klebsiella pneumoniae died readily after their addition to raw sewage, but they grew in sterilized sewage. The decline was not a result of abiotic stresses, and because the bacteria were able to survive in large numbers for at least 15 days in solutions containing no organic nutrients, it was not a result of competition. Toxin production, bacteriophages, and Bdellovibrio sp. did not cause the disappearance of the two bacterial species. A decline was also evident if the sewage was first passed through a 3-micron (pore size) filter or treated with cycloheximide or cycloheximide plus nystatin, but protozoa developed under these conditions. Little or no decline occurred if the sewage was filtered and treated with the eucaryotic inhibitors before the addition of S. typhimurium or K. pneumoniae, and protozoa were not detected. S. typhimurium increased in abundance if cycloheximide, streptomycin, and erythromycin or large amounts of glucose were added to sewage. Tetrahymena thermophilus did not significantly reduce the population of S. typhimurium in buffer when the density of the bacterium was about 10(4)/ml. However, when more than 10(8) Enterobacter agglomerans cells per ml were added to the buffer, T. thermophilus reduced the abundance of E. agglomerans and S. typhimurium to 10(6) and 10/ml, respectively. The density of S. typhimurium was further decreased by a second increment of E. agglomerans cells. The disappearance of S. typhimurium and K. pneumoniae from sewage thus is the result of predation by protozoa.(ABSTRACT TRUNCATED AT 250 WORDS)     

Orientation and food search behaviour of a deep sea lobster in turbulent versus laminar odour plumes

New Zealand scampi (Metanephrops challengeri) is a commercially important deep-water lobster species that is caught by bottom trawling on areas of muddy seafloor on the continental shelf below 300 m. Efforts are being made to develop lower impact potting methods to harvest scampi, however, they can only be caught when out of their burrows and searching for food. This emergent food searching behaviour appears to be associated with periods of higher tidal flow. Such water flow will increase turbulence along the sea floor, which has been observed to improve the efficiency of chemosensory food searching in some lobster species. Consequently, this study examined the food search behaviour of scampi in response to odours from two types of bait (mackerel and mussel) in both turbulent and laminar flows. Scampi were more efficient at foraging in the turbulent flow than in the laminar flow, using shorter search paths in response to both types of bait. Scampi in the turbulent flow reached the mussel bait 44% faster and with lower mean heading angles than in laminar flow. However, there was no difference between the flow regimes for the mackerel bait. The pattern of orientation behaviour was similar under both flow regimes, suggesting that the scampi were using the same orientation strategy, but it was more accurate in turbulent flows. The results show that the foraging efficiency of scampi improves in turbulent conditions and that this may explain their increased emergent behaviour during periods of higher tidal flows.     

Arginine metabolism in the deep sea tube worm Riftia pachyptila and its bacterial endosymbiont

The present study describes the distribution and properties of enzymes involved in arginine metabolism in Riftia pachyptila, a tubeworm living around deep sea hydrothermal vents and known to be engaged in a highly specific symbiotic association with a bacterium. The results obtained show that the arginine biosynthetic enzymes, carbamyl phosphate synthetase, ornithine transcarbamylase, and argininosuccinate synthetase are present in all of the tissues of the worm and in the bacteria. Thus, Riftia and its bacterial endosymbiont can assimilate nitrogen and carbon via this arginine biosynthetic pathway. The kinetic properties of ornithine transcarbamylase strongly suggest that neither Riftia nor the bacteria possess the catabolic form of this enzyme belonging to the arginine deiminase pathway, the absence of this pathway being confirmed by the lack of arginine deiminase activity. Arginine decarboxylase and ornithine decarboxylase are involved in the biosynthesis of polyamines such as putrescine and agmatine. These activities are present in the trophosome, the symbiont-harboring tissue, and are higher in the isolated bacteria than in the trophosome, indicating that these enzymes are of bacterial origin. This finding indicates that Riftia is dependent on its bacterial endosymbiont for the biosynthesis of polyamines that are important for its metabolism and physiology. These results emphasize a particular organization of the arginine metabolism and the exchanges of metabolites between the two partners of this symbiosis.     

Novel polymorphic microsatellite loci for a new target species,the sea cucumber Holothuria mammata

With traditional finfish fisheries declining and pushing transition to new invertebrates target species, sea cucumbers have been heavily explored, suffering global overexploitation and worldwide depletion of their stocks.Nowadays, holothurians from the Mediterranean Sea and NE Atlantic Ocean are being exported to Asian markets. The scarce knowledge about their biology, population dynamics, ecology and genetics, is promoting defective management of their fisheries.We report the development of 9 novel polymorphic microsatellites markers for Holothuria mammata and characterized them by testing in three different sample locations. All nine microsatellites revealed high polymorphism and diversity, with high number of alleles, ranged from 11 to 22 and expected heterozygosity, between 0.52 and 0.92. Significant genetic differentiation was found between populations.These microsatellites are providing valuable information which could be applied to fisheries management including, identification of stocks, assessment of their genetic diversity, estimation of gene flow and monitoring the fishery effects on exploited populations.     

Anadromous species and the new international law of the sea

Abstract

This article examines the evolution of international law relating to anadromous species, focusing exclusively on salmon and primarily considering the period since World War II. The discussion concerns major international harvesting in the western North Pacific, eastern North Pacific, and the North Atlantic. Unilateral actions are also described. Special attention is given to the relevant articles of the 1982 Convention on the Law of the Sea and to developments since its conclusion. Evidence is assessed for considering that the customary international law of the sea now recognizes the authority of the state of origin of salmon to prohibit high seas harvesting of salmon.     

Macroalgal farming in the sea: water motion and nitrate uptake

A better understanding of water motion effects on nutrient uptake by marine crop plants should make it possible to farm the sea more effectively. Farms in China, Japan and the Philippines now grow plants on slack lines or nets that move with passing waves and currents. Nutrient uptake rates are increased onLaminaria farms in China by adding nitrogen-containing fertilizer. In contrast, forests of the giant kelp,Macrocystis grow in California at low nutrient levels without fertilization. The giant kelp, compared as a structure with the slack Chinese farms, has float-supported, spring-like stipes that stretch and recoil as waves pass. This motion seems likely to enhance flow over the thallus surface. In thus study we modified flow around kelp blades in a water tunnel in the laboratory by changing orifice plates, and flow around Chinese-style long-line farms in the sea by tightening them under various sea conditions. Our measurements suggest that if marine farms were designed and operated to increase water movement over the plants being grown, their rates of nutrient uptake, and growth would increase.This paper was presented at the Symposium on Applied Phycology at the Fourth International Phycological Congress, Duke University.