The role of copepods in the planktonic ecosystems of mixed and stratified waters of the European shelf seas

The European shelf seas can be divided into regions which have tidally mixed waters and thermally stratified waters. The tidally mixed near shore environments support zooplankton communities dominated by smaller copepods and having large meroplankton contributions. These small copepods (Centropages spp., Temora spp., Acartia spp., Paral Pseudo/Microcalanus spp.) together with the microzooplankton component form a different and more complex food web than the larger copepod/diatom link associated with thermally stratified waters. The copepods Calanus finmarchicus and C. helgolandicus account for over 90% of the copepod dry weight biomass in stratified waters. Although occurring in lower numbers in mixed waters they can still make significant contributions to the biomass. A 31 year time series from the European shelf shows the inter- and intea-annual variability of these species. The basic biology and food web that these two systems support, and the transfer of energy, can result in marked differences in quantity and quality of particulates available as food for fish larvae. Calanus dominated systems allow the primary production to be directed straight through the trophic food chain (diatoms/Calanus/fish larvae) while the near shore communities of smaller copepods limit the amount of energy being transferred to the higher trophic levels. Eighty-two Longhurst Hardy Plankton Recorder hauls were used as the data base for this study. In all cases the zooplankton was dominated by copepods both in numbers and biomass accounting for > 80% of total zooplankton dry weight in the Irish Sea, Celtic Sea, shelf edge of the Celtic Sea and the northern and southern North Sea in Spring.     

Southeastern U.S.A. continental shelf respiratory rates revisited

Respiratory rates on the U. S. southeastern continental shelf have been estimated several times by different investigators, most recently by Jiang et al. (Biogeochemistry 98:101–113, 2010) who report lower mean rates than were found in earlier work and attribute the differences to analytical error in all methods used in earlier studies. The differences are, instead, attributable to the differences in the geographical scope of the studies. The lower estimates of regional organic carbon flux of Jiang et al. (Biogeochemistry 98:101–113, 2010) are a consequence of their extrapolation of data from a small portion of the shelf to the entire South Atlantic Bight. This comment examines the methodologies used as well as the variability of respiratory rates in this region over space and time.     

Sedimentary environments of the Central Region of the Great Barrier Reef of Australia

The sediments and calcareous organisms on the outer reefal shelf of the Central Region of the Great Barrier Reef were collected and observed by SCUBA diving and research vessel techniques (including underwater television) to understand the production and processes of deposition of the sediment. The carbonate grains are mainly sand and gravel size and solely of skeletal origin. Over the whole area the major CaCO₃ producers, in order of decreasing importance are: benthic foraminiferans (chiefly Operculina, Amphistegina, Marginopora, Alveolinella and Cycloclypeus), the calcareous green alga Halimeda, molluscs and corals. Coral abundance is high only close to reefs and submerged rocky substrates. Benthic foraminiferal sands dominate the inter-reef areas i.e. the bulk of the shelf, and Halimeda gravels form an outer shelf band between 60 and 100 m depths. Seven distinct facies are recognised after quantitative analyses of the sediments. These are: A. Shelf edge slope (>120 m depth); B. Shelf edge (with rocky outcrops); C. Outer shelf with high Halimeda (>40%); D. Inter-reef I; E. Inter-reef II ( 100 m depth but >2% pelagics); F. Lee-ward reef talus wedge (<2 km from sea level reefs); G. Lagoonal.     

Bryozoans of the Weddell Sea continental shelf,slope and abyss: did marine life colonize the Antarctic shelf from deep water,outlying islands or in situ refugia following glaciations?

Aim At the height of glaciations such as the Last Glacial Maximum (LGM), benthic life on polar continental shelves was bulldozed off nearly all of the Antarctic shelf by grounded ice sheets. The origins of the current shelf benthos have become a subject of considerable debate. There are several possible sources for the current Antarctic shelf fauna, the first of which is the continental slope and deep sea of the Southern Ocean. The high levels of reported eurybathy for many Antarctic species are taken as evidence supporting this. A second possible source for colonists is the southern margins of other continents. Finally, shelves could have been recolonized from refugia on the continental shelves or slopes around Antarctica. The current study investigates whether the patchily rich and abundant biota that now occurs on the Antarctic continental shelf recolonized from refugia in situ or elsewhere. Location Weddell Sea, Antarctica. Methods We examined bryozoan samples of the BENDEX, ANDEEP III and SYSTCO expeditions, as well as the literature. Using similarity matrices (Sørensen coefficient), we assessed similarities of benthos sampled from around Antarctica. By assessing numbers of species shared between differing depths and adjacent shelf areas, we evaluated the origins of cheilostome bryozoan communities. Results Bryozoans decreased from 28, 6.5 and 0.3 colonies per trawl, and 0.16, 0.046 and 0.0026 colonies per cm² of hard surface from shelf to slope to abyssal depths. We found little and no support for recolonization of the Weddell Sea shelf by bryozoans from the adjacent slope and abyss, in the scenario of LGM faunal wipe‐out. The Weddell Sea shelf bryozoan fauna was considerably more similar to those on other Antarctic shelves than to that of the adjacent (Weddell Sea) continental slope. The known bryozoan fauna of the Weddell Sea shelf is not a subset of the Weddell Sea slope or abyssal faunas. Main conclusions We consider that the composition of the current Weddell Sea bryozoan fauna is most easily explained by in situ survival. Thus we consider that at least some of the Weddell Sea fauna persisted throughout the LGM, although not necessarily at the same locations throughout, to recolonize the large area currently occupied.     

Environmental distribution of colony growth form in the favositid Pleurodictyum americanum

Pandolfi, John M. & Burke, Collette D. 1989 01 15: Environmental distribution of colony growth form in the favositid Pleurodictyum americanum. Lethaia , Vol. 22, pp. 69–84. Oslo. ISSN 0024–1164.
Colony growth form is of fundamental importance to understanding the ecology of both modern and ancient marine sessile colonial animals. Fourier shape analysis of the coral Pleurodictyun americantum (Tabulata: Favositida) from the Middle Devonian Hamilton Group of New York State, indicates that colony growth form is variable between environments. Discriminate function analysis of harmonics 2–6 of 51 assemblages of Pleurodictyum americanum (N = 1900) shows that this species displays an onshore to offshore gradient in colony shape. Offshore environments characterized by low levels of turbidity, oxygen, and light contain more flattened, less hemispherical growth forms, whereas onshore environments characterized by high levels of turbidity, oxygen, and light contain more hemispherical, less flattened growth forms. Harmonic shape analysis detected subtle differences among samples of P. americanum from different environments, but also showed that distinctive morphotypes are distributed within horizons. as well as between them. In fact, no one-to-one correspondence in growth form to environment is apparent; growth forms are distributed within environments, suggesting that genetic factors may have had a greater influence over coral growth form than environment. In tabulate corals, patterns of within species variability must be determined before growth form may be useful in interpreting ancient environments. □ Tabulate coral, Fourier shape analysis, morphological variability, growth form, Devonian, Hamilton Group, New York .     

Individual foraging site fidelity in lactating New Zealand fur seals: Continental shelf vs. oceanic habitats

Wide‐ranging marine central place foragers often exhibit foraging site fidelity to oceanographic features over differing spatial scales (i.e., localized coastal upwellings and oceanic fronts). Few studies have tested how the degree of site fidelity to foraging areas varies in relation to the type of ocean features used. In order to determine how foraging site fidelity varied between continental shelf and oceanic foraging habitats, 31 lactating New Zealand fur seals (Arctocephalus australis forsteri 1 ) were satellite tracked over consecutive foraging trips (14–108 d). Thirty‐seven foraging trips were recorded from 11 females that foraged on the continental shelf, in a region associated with a coastal upwelling, while 65 foraging trips were recorded from 20 females that foraged in oceanic waters. There were no significant differences in the mean bearings (to maximum distance) of individual's consecutive foraging trips, suggesting individual fidelity to foraging areas. However, overlap in area and time spent in area varied considerably between continental shelf and oceanic foragers. Females that foraged on the continental shelf had significantly greater overlap in consecutive foraging trips when compared to females that foraged in oceanic waters (overlap in 5 × 5 km grid cells visited on consecutive trips 55.9%± 20.4% and 13.4%± 7.6%, respectively). Females that foraged on the continental shelf also spent significantly more time within the same grid cell than females that foraged in oceanic waters (maximum time spent in 5 × 5 km grid cells: 14%± 5% and 4%± 2%, respectively). This comparatively high foraging site fidelity may reflect the concentration of productivity associated with a coastal upwelling system, the Bonney Upwelling. Lower foraging site fidelity recorded by seals that foraged in oceanic waters implies a lower density/larger scale habitat, where prey are more dispersed or less predictable at fine scales, when compared to the continental shelf region.     

Seasonal patterns of winter flounder Pseudopleuronectes americanus abundance and reproductive condition on the New York Bight continental shelf

To resolve varied and sometimes conflicting accounts of spawning and habitat characteristics for winter flounder Pseudopleuronectes americanus, seasonal patterns in abundance and reproductive condition were investigated in the New York Bight, near the southern edge of their current reproductive range. Fish were collected from trawl surveys on the inner continental shelf from October 2006 to October 2007. Pseudopleuronectes americanus were most abundant during January and April surveys, were rarely collected in August, with intermediate abundances in June and October. Measurements of fish condition [hepato‐somatic index (I_H), condition factor (K) and the per cent dry mass of muscle tissue (%M_D)] and reproductive condition [gonado‐somatic index (I_G)] were determined to evaluate seasonal changes in energy accumulation and depletion and reproduction. Males and females had similar patterns in body and reproductive condition, although the magnitude of change was greater for females. I_H values were highest during spring and early summer, suggesting increased feeding following spawning. K and %M_D increased through spring and summer then declined in the autumn and winter concurrent with gonadal development. Gonads began developing in the autumn, and in January, I_G values approached spawning levels, with many spent individuals collected in spring. Within these general patterns, however, there was a large degree of variability among individuals, and a few mature non‐reproductive (‘skipped spawning’) females were observed. In the period after spawning, increased energy intake, indicated by increased I_H, may influence reproductive output since this energy is gradually transferred to the muscle and used for gonadal development in the forthcoming year. The occurrence of ripening individuals on the inner continental shelf in January suggests that these fish either rapidly move into estuaries to spawn by February–March or they remain on the inner shelf to spawn, or some combination of these. Future studies should evaluate these possibilities, as both estuarine and inner shelf habitats are potentially affected by activities such as dredging, sand dredging and wind energy development.     

古尼拟青霉表型多样性及其与活性成分的关系

目的研究表型多样性与活性成分之间的联系。方法研究10株古尼拟青霉菌株的培养特性,观察菌落形态,测定其产孢量和粗蛋白、多糖、甘露醇、麦角甾醇、腺苷等主要活性成分含量。结果A类菌株菌落背面褶皱不平,中央为棕色环状,周边为淡黄色,是生长性状优良、活性成分含量高,适于大规模工业化生产的优质菌株。结论不同菌株具有不同的形态特征,且形态特征与其产孢量及主要活性成分存在一定内在联系。     

Quantifying coral morphology

Coral morphology has important implications across scales, from differences in physiology, to the environments they are found, through to their role as ecosystem engineers. However, quantifying morphology across taxa is difficult, and so morphological variation is typically captured via coarse growth form categories (e.g. arborescent and massive). In this study, we develop an approach for quantifying coral morphology by identifying continuous three-dimensional shape variables. To do so, we contrast six variables estimated from 152 laser scans of coral colonies that ranged across seven growth form categories and three orders of magnitude of size. We found that 88% of the variation in shape was captured by two principal components. The main component was variation in volume compactness (cf. convexity), and the second component was a trade-off between surface complexity and top-heaviness. Variation in volume compactness also limited variation along the second axis, where surface complexity and top-heaviness ranged more freely when compactness was low. Traditional growth form categories occupied distinct regions within this morphospace; however, these regions overlapped due to scaling of shape variables with colony size. Nonetheless, with four of the shape variables we were able to predict traditional growth form categories with 70 to 95% accuracy, suggesting that the continuous variables captured most of the qualitative variations implied by these growth forms. Distilling coral morphology into continuous variables that capture shape variation will allow for better tests of the mechanisms that govern coral biology, ecology and ecosystem services such as reef building and provision of habitat.

     

The role of continental shelf width in determining freshwater phylogeographic patterns in south‐eastern Australian pygmy perches (Teleostei: Percichthyidae)

Biogeographic patterns displayed by obligate freshwater organisms are intimately related to the nature and extent of connectivity between suitable habitats. Two of the more significant barriers to freshwater connections are seawater and major drainage divides. South‐eastern Australia provides a contrast between these barriers as it has discrete areas that are likely influenced to a greater or lesser extent by each barrier type. We use continental shelf width as a proxy for the potential degree of river coalescence during low sea levels. Our specific hypothesis is that the degree of phylogeographic divergence between coastal river basins should correspond to the continental shelf width of each region. This predicts that genetic divergences between river basins should be lowest in regions with a wider continental shelf and that regions with similar continental shelf width should have similar genetic divergences. Pygmy perches (Nannoperca australis and Nannoperca ‘flindersi’) in south‐eastern Australia provide an ideal opportunity to test these biogeographic hypotheses. Phylogeographic patterns were examined based on range‐wide sampling of 82 populations for cytochrome b and 23 polymorphic allozyme loci. Our results recovered only limited support for our continental shelf width hypothesis, although patterns within Bass clade were largely congruent with reconstructed low sea‐level drainage patterns. In addition, we identified several instances of drainage divide crossings, typically associated with low elevational differences. Our results demonstrate high levels of genetic heterogeneity with important conservation implications, especially for declining populations in the Murray–Darling Basin and a highly restricted disjunct population in Ansons River, Tasmania.     

Evaluation of the apron as a taxonomic trait in ixodid ticks of the genus Dermacentor Koch (Ixodoidea, Ixodidae)

Study of the apron in 9 species of the genus Dermacentor from the fauna of the USSR has revealed differencies in its structure. The subgenus Dermacentor (s. str.) differs from two other subgenera both in the shape of the apron itself and in the shape of the postgenital sclerite and setae of perigenital area. Close species within each of two other subgenera differ in apron proportion, shape and size of denticles along its hind edge, and sometimes in their number. Inspite of the statistically reliable interspecific differences in apron structure a wide range of individual variability of some details and geographical specificity of samples from various places of the area were observed in species with a vast area.     

Topographic and faunal studies of shelf-edge prominences off the central eastern Florida coast

Eighty topographic transects were made off the central Atlantic coast of Florida between Cape Canaveral and Palm Beach, November, 1973 to September, 1974, aboard the Research Vessel Gosnold. Profiles obtained with a precision depth recorder indicated the presence of continuous and discontinuous structures on the outer continental shelf and at the shelf-slope break. Most conspicuous was a band of pinnacles, benches, mounds and troughs extending north from Fort Pierce to Cape Canaveral, Florida, and a massive mound occurring off St. Lucie Inlet. These occurred under the western edge of the Florida Current and along the edge of the continental shelf (about 80° W). From Fort Pierce southward to Palm Beach, these structures almost disappear. Dredgings in two selected areas of pronounced vertical profile, and preliminary visual and photographic observations made with the Johnson-Sea-Link I submersible, confirmed the existence of rich, sessile and motile invertebrate assemblages and fish populations associated with exposed limestone bedrock, talus, and the scleractinian coral Oculina varicosa.     

Do freshwater ecoregions and continental shelf width predict patterns of historical gene flow in the freshwater fish Poecilia butleri?

We examined historical patterns of gene flow in the freshwater fish Poecilia butleri in western Mexico. We tested the hypothesis that the boundaries between four freshwater ecological communities (ecoregions) might have limited the movement of P. butleri because changes in species compositions might restrict establishment between adjacent ecoregions, even in situations where a physical barrier is absent. Hence, we predicted that boundaries between ecoregions should correspond to phylogeographical breaks in P. butleri. We also tested the hypothesis that the width of the continental shelf affected historical gene flow in P. butleri because a broad continental shelf provides a greater opportunity for rivers to coalesce during historical episodes of low sea levels as opposed to a narrow continental shelf that should restrict the potential for gene flow among adjacent rivers. Hence, we predicted greater amounts of historical gene flow among neighbouring river basins in the region of western Mexico where the continental shelf is wider, whereas, in the region where the continental shelf is narrower, we expected to detect limited levels of historical gene flow. We analyzed mitochondrial DNA sequence data (cytochrome b) taken from 264 individuals of P. butleri collected from 34 locations distributed across four different ecoregions in western Mexico. To examine patterns of phylogenetic diversification and historical gene flow in P. butleri, we employed several analytical approaches, including traditional tree‐based phylogenetic analyses (likelihood and parsimony), haplotype network reconstruction, analyses of molecular variance, and spatial analysis of molecular variance. We found genetic breaks coinciding with two out of three different ecoregion boundaries, suggesting limited historical gene flow. In addition to different species compositions between these adjacent ecoregions, geological features such as the Trans‐Mexican Volcanic Belt and the mountainous topography in south‐western Mexico, likely contributed to these observed genetic breaks. By contrast, no genetic break was evident between two other ecoregions, a result that partially rejects our first hypothesis. Several results were consistent with our second hypothesis. Changes in the width of the continental shelf in western Mexico are associated with the observed patterns of historical gene flow. Our results indicate that the interactions among multiple geological and biological factors affect the spatial patterns of genetic diversity of widespread freshwater species. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 112 , 399–416.     

Extreme,continuous variation in an island snail: local diversification and association of shell form with the current environment

On Rosemary Island, a small continental island (11 km²) in the Dampier Archipelago, Western Australia, snails of the genus Rhagada have extremely diverse morphologies. Their shells vary remarkably in size and shape, with the latter ranging from globose to keeled‐flat, spanning the range of variation in the entire genus. Based primarily on variation in shell morphology, five distinct species are currently recognized. However, a study of 103 populations has revealed continuity of shell form within a very closely‐related group. A phylogenetic analysis of specimens from Rosemary Island, and other islands in the Dampier Archipelago, indicates that much of the morphological variation has evolved on the island, from within a monophyletic group. Within the island, snails with distinct shell morphologies could not be distinguished based on variation in mitochondrial DNA or their reproductive anatomy. The shell variation is geographically structured over a very fine scale, with clines linking the extreme forms over distances less than 200 m. Although there is no evidence that the different forms have evolved in isolation or as a consequence of drift, there is a strong association between keeled‐flat shells and rocky habitats, suggesting that shell shape may be of adaptive significance. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 756–769.     

Species diversity and distribution within the deep-sea nematode genus Acantholaimus on the continental shelf and slope in Antarctica

Species diversity and distribution—in relation with biogeography, bathymetry and environmental characteristics—within the genus Acantholaimus were investigated on the shelf and slope (182–2,009 m) in the Weddell Sea (Kapp Norvegia and Vestkapp) and near the Antarctic Peninsula (Drake Passage and Bransfield Strait). The occurrence of the otherwise typical deep-sea genus Acantholaimus in high densities and diversity on the continental shelf is a unique feature of the Southern Ocean. Fifty-five Acantholaimus species (morpho-types) were distinguished in our study area, of which 39 were found on the shelf. The Acantholaimus communities were characterised by a high number of congeneric species and the presence of many rare species, similar to typical deep-sea communities. Fourteen species had a distribution extending from the shelf to the lower slope. High local and regional diversity was obvious and a considerable level of endemism was suggested. A negative impact of chlorophyll a concentrations, reflecting the availability of phytoplankton, was found on the Acantholaimus densities and species diversity.     

东海大陆架鱼类群落的空间结构

根据1997～2000年东海大陆架水深30～200米海域4个航次的底拖网调查资料,运用多元分析(聚类分析和非度量多维标度(NMDS))、SIMPER以及BIO-ENV方法分析了东海鱼类群落的空间格局以及与非生物环境因子之间的关系。结果表明,东海大陆架海域鱼类群落在空间上可分为3个群落类型,分别为东海近海群落、东海大陆架外缘群落以及这两个群落之间的东海大陆架混合群落。东海大陆架鱼类群落的种类组成沿着水深梯度的变化明显,每种鱼类均有一定的水深范围。不同的流系具有不同的温盐特征,影响了鱼类的分布及其洄游,从而控制了不同鱼类群落类型的种类组成。东海大陆架鱼类群落类型中,大陆架外缘群落类型各调查季节在空间分布和种类组成上相对稳定,而东海近海群落类型和东海大陆架混合群落类型在夏季融合在一起,其它季节保持各自的空间结构。同时东海陆架区只在一种群落类型中出现的鱼类相对较少,多数鱼类同时出现在两个群落类型中或3个群落类型中,但是它们相对较集中的区域明显不同,在不同群落类型中的生物量和出现频率有很大的差异,这些鱼类在其不同的生长发育阶段由于对海洋环境的不同需求,它们根据繁殖或摄食需求而在东海陆架区范围内洄游以寻找最适的海洋环境,因此对3个群落类型定义是相对的,很难严格确定一个永久的鱼类群落地理区域,以形成相互隔离的鱼类群落,而且鱼类群落之间的边界随着季节不同发生变化。     

Taking a trip to the shelf: Behavioral decisions are mediated by the proximity to foraging habitats in the black‐legged kittiwake

For marine top predators like seabirds, the oceans represent a multitude of habitats regarding oceanographic conditions and food availability. Worldwide, these marine habitats are being altered by changes in climate and increased anthropogenic impact. This is causing a growing concern on how seabird populations might adapt to these changes. Understanding how seabird populations respond to fluctuating environmental conditions and to what extent behavioral flexibility can buffer variations in food availability can help predict how seabirds may cope with changes in the marine environment. Such knowledge is important to implement proper long‐term conservation measures intended to protect marine predators. We explored behavioral flexibility in choice of foraging habitat of chick‐rearing black‐legged kittiwakes Rissa tridactyla during multiple years. By comparing foraging behavior of individuals from two colonies with large differences in oceanographic conditions and distances to predictable feeding areas at the Norwegian shelf break, we investigated how foraging decisions are related to intrinsic and extrinsic factors. We found that proximity to the shelf break determined which factors drove the decision to forage there. At the colony near the shelf break, time of departure from the colony and wind speed were most important in driving the choice of habitat. At the colony farther from the shelf break, the decision to forage there was driven by adult body condition. Birds furthermore adjusted foraging behavior metrics according to time of the day, weather conditions, body condition, and the age of the chicks. The study shows that kittiwakes have high degree of flexibility in their behavioral response to a variable marine environment, which might help them buffer changes in prey distribution around the colonies. The flexibility is, however, dependent on the availability of foraging habitats near the colony.     

Estimating denitrification in North Atlantic continental shelf sediments

A model of coupled nitrification/denitrification was developed for continental shelf sediments to estimate the spatial distribution of denitrification throughout shelf regions in the North Atlantic basin. Using data from a wide range of continental shelf regions, we found a linear relationship between denitrification and sediment oxygen uptake. This relationship was applied to specific continental shelf regions by combining it with a second regression relating sediment oxygen uptake to primary production in the overlying water. The combined equation was: denitrification (mmol N m^–2 d^–1)=0.019^* phytoplankton production (mmol C m^–2 d^–1). This relationship suggests that approximately 13% of the N incorporated into phytoplankton in shelf waters is eventually denitrified in the sediments via coupled nitrification/denitrification, assuming a C:N ratio of 6.625:1 for phytoplankton. The model calculated denitrification rates compare favorably with rates reported for several shelf regions in the North Atlantic.The model-predicted average denitrification rate for continental shelf sediments in the North Atlantic Basin is 0.69 mmol N m^{– 2} d^–1. Denitrification rates (per unit area) predicted by the model are highest for the continental shelf region in the western North Atlantic between Cape Hatteras and South Florida and lowest for Hudson Bay, the Baffin Island region, and Greenland. Within latitudinal belts, average denitrification rates were lowest in the high latitudes, intermediate in the tropics and highest in the mid-latitudes. Although denitrification rates per unit area are lowest in the high latitudes, the total N removal by denitrification (53 × 10¹⁰ mol N y^–1) is similar to that in the mid-latitudes (60 × 10¹⁰ mol N y^–1) due to the large area of continental shelf in the high latitudes. The Gulf of St. Lawrence/Grand Banks area and the North Sea are responsible for seventy-five percent of the denitrification in the high latitude region. N removal by denitrification in the western North Atlantic (96 × 10¹⁰ mol N y^–1) is two times greater than in the eastern North Atlantic (47 × 10¹⁰ mol N y^–1). This is primarily due to differences in the area of continental shelf in the two regions, as the average denitrification rate per unit area is similar in the western and eastern North Atlantic.We calculate that a total of 143 × 10¹⁰ mol N y^–1 is removed via coupled nitrification/denitrification on the North Atlantic continental shelf. This estimate is expected to underestimate total sediment denitrification because it does not include direct denitrification of nitrate from the overlying water. The rate of coupled nitrification/denitrification calculated is greater than the nitrogen inputs from atmospheric deposition and river sources combined, and suggests that onwelling of nutrient rich slope water is a major source of N for denitrification in shelf regions. For the two regions where N inputs to a shelf region from onwelling have been measured, onwelling appears to be able to balance the denitrification loss.     

Colonial and Cellular Polymorphism in Xenorhabdus luminescens

A highly polymorphic Xenorhabdus luminescens strain was isolated. The primary form of X. luminescens was luminescent and nonswarming and produced a yellow pigment and antimicrobial substances. The primary form generated a secondary form that had a distinct orange pigmentation, was weakly luminescent, and did not produce antimicrobial substances. Both the primary and secondary forms generated a set of colony variants at frequencies that exceeded normal rates for spontaneous mutation. The variant forms include nonswarming and swarming forms that formed large colonies and a small-colony (SC) form. The primary and secondary forms generated their SC forms at frequencies of between 1 and 14% and 1 and 2%, respectively. The SC forms were distinct from their parental primary and secondary forms in colony and cellular morphology and in protein composition. The cellular morphology and protein patterns of the nonswarming and swarming colony variants were all very similar. The DNA fingerprints of all forms were similar. Each SC-form colony reverted at high frequency to the form from which it was derived. The proportion of parental-type cells in the SC-form colonies varied with age, with young colonies containing as few as 0.0002% parental-type cells. The primary-to-secondary switch was stable, but all the other colony forms were able to switch at high frequencies to the alternative colony phenotypes.