Blooms of tunicates Oikopleura spp. and Dolioletta gegenbauri in the Seto Inland Sea, Japan, during summer

The ecological importance of appendicularians Oikopleura dioica and O. longicauda, and a doliolid Dolioletta gegenbauri as metazoan secondary producers was assessed in summer 1997 in the Seto Inland Sea, Japan. Blooms by the appendicularians occurred twice within a month following peaks of pico-/nanoplankton biomass. The biomass of Oikopleura spp. averaged over the water column (<B> ranged 0.1 to 8.0 μgC l-¹ and temporal average of <B> over the survey period ( ) was 3.2 μgC l-¹, 70% of for calanoid copepods. Furthermore, the temporal average of the production for Oikopleura spp. was 2.4 times higher than that for calanoids, reflecting the extraordinary high growth rates of the appendicularians. D. gegenbauri developed its population following a diatom bloom and <B> reached to 37 μgC l-¹ at the peak time. Although individuals of D. gegenbauri were seldom observed during the first half of the survey, for the doliolid (8.4 μgC l-¹) nearly doubled that for calanoids. These results indicate that the gelatinous tunicates Oikopleura spp. and D. gegenbauri play important roles as metazoan secondary producers in the Seto Inland Sea during summer. This revised version was published online in September 2006 with corrections to the Cover Date.     

Living below the halocline: strategies of deep-living species in the highly stratified brackish Bornholm Basin (central Baltic Basin)

The Baltic Sea is the largest brackish water area of the world.On the basis of the data from 16 cruises, we show the seasonaland vertical distribution patterns of the appendicularians Fritillariaborealis, Oikopleura dioica and the cyclopoid copepod Oithonasimilis, in the highly stratified Bornholm Basin. These specieslive at least temporarily below the permanent halocline anduse different life strategies to cope with the brackish environment.The cold-water species F. borealis is abundant in the upperlayers of the water column before the thermocline develops.With the formation of the thermocline abundance decreases andthe specimens outlast higher temperatures below the halocline.Distribution and strategy suggest that F. borealis might bea glacial relict species in the Baltic Sea. Although Oikopleuradioica is only abundant during summer, O. similis is presentall year round. Both species have in common that their verticaldistribution is restricted to the waters below the halocline,most likely due to their requirements of higher salinities.We argue that the observed strategies are determined by ecophysiologicalconstraints and life history traits. These species share anomnivorous feeding behaviour and the capability to utilise aspectra of small particles as food. As phytoplankton concentrationis negligible below the halocline, we suggest that these speciesfeed on organic material and heterotrophic organisms that accumulatein the density gradient of the halocline. Therefore, the deephaline waters in the Baltic Sea represent a habitat providingshelter from predation and food supply for adapted species thatallows them to gather sufficient resources and to maintain populations.     

Appendicularian assemblages in a shelf area and their relationship with temperature

The structure of the community of appendicularians was describedby multivariate analyses throughout a seasonal cycle on thecentral Cantabrian coast. It is shown by correlation and principalcomponents analysis that the appendicularian species may bearranged in a successional sequence in relative abundance thatis closely coupled to a temperature gradient. This sequencestarts with Fritillaria borealis, which exhibits highest relativeabundance during January, being sequentially followed by Oikopleuradioica, Fritillaria pellucida, Oikopleura fusiformis and Oikopleuralongicauda. This species numerically dominated the communityfrom September to December. Sea surface temperature and thetemperature at the depth of the chlorophyll maximum were bothreliable predictors of the species composition of the community.However, only the latter provided an adequate explanation forthe persistence of cryophilic communities in stratified oceanicenvironments and the dominance of thermophilic communities afterthe autumn mixing period. Under stratified conditions, surfacetemperatures are high (up to 21°C), but temperatures atthe depth of the chlorophyll maximum are low (<15°C).These differences disappear after the autumn mixing, when thewater column exhibits a uniform temperature profile (16°C).Critically, however, although there is a sharp decline in surfacetemperature, water at the depth of the chlorophyll maximum iswarmer than during stratification. A simple conceptual modelis proposed to account for these features and predictions aremade regarding the vertical distribution of appendiculariansduring stratification. The relevance of non-anthropomorphictemperature measures, such as the temperature at the depth ofthe chlorophyll maximum, for other zooplankton groups is alsodiscussed.     

Zooplankton growth rates: the larvaceans Appendicularia, Fritillaria and Oikopleura in tropical waters

The growth rates of Appendicularia sicula, Fritillaria borealissargassi, Fritillaria haplostoma, Oikopleura dioica and Oikopleuralongicauda were determined from microcosms incubated in situat 23C in Jamaican waters. Experiments were conducted fromoligotrophic offshore waters, through mesotrophic Lime Cay andeutrophic Kingston Harbour in both natural and nutrient-enhancedphytoplankton communities. Length-weight relationships werecalculated for two of these species: O.longicauda log W=2.47log TL –6.10 and F.haplostoma log W=2.44 log TL –7.37,where weight (W) is in micrograms and trunk length (TL) is inmicrometres. Instantaneous growth rates averaged 1.7–2.5day^–1 for the five species and were observed as high as3.3 day^–1 These instantaneous rates are equivalent todaily specific growth rates averaging 4.6–11.4 and rangingup to 28. In larger genera, growth rates were related positivelyto picoplankton and nanoplankton concentration, and negativelyto the biomass of larvaceans, but in the smallest species growthwas unrelated to these factors. However, because the variabilityin these two factors within microcosms exceeded their naturalrange of variability, growth rates of larvaceans may normallybe unlimited by resources or population density effects. ¹Present address :Monterey Bay Aquarium Research Institute 7700Sandholdt Road, Moss Landing, CA 95039-0628, USA ²Present address :Bedford Institute of Oceanography PO Box 1006,Dartmouth, Nova Scotia B2Y 4A2, Canada     

Appendicularians: an important food supply for the Argentine anchovy Engraulis anchoita in coastal waters

The stomach contents of 532 adult specimens of the Argentine anchovy, Engraulis anchoita, caught in coastal waters of the Argentine Sea from 1994 to 1996, were analyzed. Larger amounts of food were found in the stomachs of anchovies collected in the northern surveys (35–40°S), with the highest values attained in 1994. Main diet items were copepods, appendicularians, cladocerans, fish eggs, and pteropods. Anchovies caught in the southern surveys (40–45°S) ingested less food, whereby the main items were copepods and appendicularians. Plankton samples collected simultaneously with the fishing trawls were also analyzed. Except for Fritillaria borealis, which occurred only in plankton samples, the same appendicularian species (Oikopleura dioica and O. fusiformis) were found in both stomach contents and in plankton samples. The Ivlev selectivity index calculated for O. dioica and O. fusiformis and for the different maturity stages of both species were in all cases ～0, supporting the hypothesis of a non‐selective feeding.     

Cellulose in the house of the appendicularianOikopleura rufescens

Summary By electron diffraction analysis, highly crystalline cellulose Iβ was found in the house (a special structure in which the tunicate lives) of the appendicularianOikopleura rufescens. Cellulose microfibrils 20 nm in width were observed in a random array or highly organized with rectangular spacing of 2 to 10 (im in the house. The bundled cellulose microfibrils formed in the inlet filters, which are highly ordered meshwork structures. This paper provides the first account of the existence of cellulose in the house of an appendicularian. Our findings showed that the house and tunic are homologous tissues among the tunicates, and that the common ancestor of the tunicates (ascidians, thaliaceans, and appendicularians) already possessed cellulose-biosynthetic ability.     

Ontogeny of the appendicularian <Emphasis Type="Italic">Oikopleura dioica</Emphasis> (Tunicata,Chordata) reveals characters similar to ascidian larvae with sessile adults

Appendicularians have always occupied a central role in considerations of tunicate and chordate evolution. Two hypotheses have been proposed – one holds that appendicularia represents the sister taxon to the remaining tunicates, the other suggests that appendicularians were derived from an ascidian-like ancestor. In the present study I report results from electron microscopic investigation of larval tunicates including the first electron microscopic investigation of the tail of the early ontogenetic appendicularian “Streckform” and discuss their phylogenetic implications. The early “Streckform” of Oikopleura dioica Fol, 1872 is invested with an extracellular covering that consists of an inner electron-light layer and an electron-dense outermost layer. In addition, the extracellular covering forms fin blades. Because these traits are shown to be similar to the tunic of different ascidian larvae, the extracellular covering in early appendicularian embryos is suggested to be homologous to the larval tunic of ascidian larvae. Overall, the tail of early developmental stages of appendicularians consists of a mosaic of apomorphic and plesiomorphic features. The straight, continuous endodermal strand was inherited from a common chordate ancestor whereas the finlets of larvae, consisting of extracellular material, were inherited from a common tunicate ancestor. The horizontal orientation of the tail as a whole was inherited from the last common ancestor of appendicularians and aplousobranch ascidians, and the discovered floating extension at the posterior tip of the tail is unique to the holoplanktonic Oikopleura dioica. These findings support the hypothesis that Appendicularia is derived from a sessile, ascidian-like ancestor.     

Production of Oikopleura longicauda (Tunicata: Appendicularia) in Toyama Bay, southern Japan Sea

Oikopleura longicauda occurred throughout the year in ToyamaBay, southern Japan Sea, and analysis of its size compositionand maturity revealed that reproduction was continuous overtheyear. Somatic growth production (P_g) varied with season from0.03 to 103 mg carbon (C) m^–2day^–1 (annual P_g 4.5g C m^–2), and house production (P_e) from 0.11 to 266 mgC m^–2 day^–1 (annualP_e 11.3 g C m^–2). The annualP_g/B ratio was 176. Compared with production data of some predominantzooplankton species in Toyama Bay, it is suggested that despitetheir smaller biomass, appendicularians are an important secondaryproducer.     

RAPD technique for taxonomic studies of Pellia epiphylla‐complex (Hepaticae, Metzgeriales)

The random amplified polymorphic DNA (RAPD) technique was used to study the genomic relationships of three liverworts from the genus Pellia: P. epiphylla, P. borealis and P. neesiana. Altogether 150 characters (150 DNA fragments obtained using PCR) were scored. These characters were used to create a matrix of pairwise distances between all the pairs of taxa. Both distance (UPGMA, Fitch–Margoliash and Neighbor–Joining) and binary character‐state (Wagner and Camin‐Sokal parsimony and compatibility) methods were applied for trees' construction. Our results strongly support distinction of the recently discovered sibling species of P. epiphylla – species N and P. epiphylla – species S, which have an allopatric distribution in Poland (N – North, S – South Poland). Moreover, our data also supports the hypothesis of a hybrid origin (alloploid) of the polyploid P. borealis from P.epiphylla−N×P.epiphylla−S. P. neesiana was excluded as a donor of either of the genomes of P. borealis. This revised version was published online in July 2006 with corrections to the Cover Date.     

Observations on the vertical distribution of zooplankton, including post-larval teleosts, off Plymouth in the presence of a thermocline and a chlorophyll-dense layer

During 1973 – 1977, when Sagitta elegans was returningto dominance over S. setosa in Plymouth waters, there was often,in summer in stratified conditions, a community characterizedby S. setosa in the warm water above the thermocline, and atypical S. elegans community in the colder layer below the thermocline.This vertical separation persisted at night even though somespecies from the S. elegans community migrated up into the warmlayer. Such vertical migration was not stopped by a temperaturediscontinuity of from 3° to 6°C, but fewer individuals,or a lesser proportion of species, took part at the higher temperaturedifference. Only a small part of the zooplankton, comprisingsmall individuals of Limacina retroversa, the appendicularians,and, possibly, post-larvae of Arnoglossus laterna, showed signsof aggregation at the thermocline or the chlorophyll maximumclose to it.     

Vertical distributions of zooplankton across the Almeria-Oran frontal zone (Mediterranean Sea)

The vertical distribution of zooplankton in the upper 700 mof the water column was investigated across the Almeria–Oranfront (Alboran Sea) during winter 1997/98 (Almofront 2 cruise).Eight sites corresponding to three hydrodynamic structures (theAtlantic jet, an anticyclonic gyre and the surrounding Mediterraneanwaters) were explored, by day and at night, using multiple-netsampling. Results are presented for the major species of copepods(>500 µm), macroplankton and microplankton. Speciesinhabiting the surface layer or occupying it at night extendeddeeper in the gyre. On the other hand, mesopelagic species (e.g.Pleuromamma borealis, Pleuromamma abdominalis, Euphausia krohniand Cyclothone braueri) were found to live deeper, at leastby day for the migrants, in the Mediterranean waters comparedwith the two other structures. These changes in preferentialdepths across the front are discussed in relation to environmentalfeatures, such as light level, abundance of food and mixed layerdepth. This study also documented the vertical distributionof some overwintering copepod species (Calanus helgolandicus,Eucalanus hyalinus and Eucalanus monachus).     

Vertical structure of small metazoan plankton, especially noncalanoid copepods. I. Deep Arabian Sea

The abundance and vertical distribution of micro-metazoans sampledwith fine nets of 0.05 mm mesh size were studied at three stationsin the Arabian Sea during the intermonsoon period (April/May1987) and down to 1850 m depth. In the epipelagic zone (0–100m). values of biomass and metazoan abundance tended to be higherthan those reported for other tropical oceanic areas. In themesopelagic zone, which is characterized by an extreme oxygendeficiency between 100 and 1000 m depth, the abundance of metazoantaxa and species numbers of non-calanoid copepods were largelyreduced. However, intermediate abundance maxima occurred withinthis zone, which were dominated by specific metazoan taxa (copepods.appendicularians) and species of non-calanoids (Oncaea sp. C).In the bathypelagic zone below 1050 m, the species diversityof the dominant copepod family Oncaeidae increased substantially.Two-thirds of a total of 69 oncaeid species recovered were confinedto this layer. As most of them were small in size and occurredin low abundance only, the increase in total oncaeid densityand/or plankton biomass was less conspicuous. Dominant Oncaeaspecies in the bathypelagic zone were O.longipes and O.brodskii.The results are compared with published data from the ArabianSea and other tropical oceanic areas with and without an extrememesopelagic oxygen minimum zone Possible causes of the intermediateabundance maxima within the oxygen deficiency zone are discussed.     

Small-scale distribution of macroplankton and micronekton in the Ligurian Sea (Mediterranean Sea) as observed from the manned submersible Cyana

A series of eight submersible dives (the MIGRAGEL I cruise)was made during late April 1986 using the French submersibleCyana to investigate macrozooptankton in the upper 400–700m of the water column. Paired day and night dives were madeat stations 3, 6, 13 and 23 nautical miles off Cape Ferrat,near Villefranche-sur-Mer, France; the distances represent differentareas in the frontal system of the Ligurian Sea. Detailed day/nightvertical distribution data are shown for the most abundant species;these include the narcomedusa Solmissus albescens, teleost fishCyclothone spp., small appendiculanans (primarily Oikopleuraalbicans), large appendicularians (an undescribed oikopleurid),diphyid siphonophores (mostly Chelophyes appendiculata) andan abundant lobate ctenophore. Salps, pyrosomes, amphipods (Phroniumasedentaria), pteropods (Cavolinia inflexa), macroscopic ‘star-like’protozoa and marine snow are also briefly discussed. The coastalzone was dominated by small appendicularians in the upper layers,with other filter feeders including large appendicularians indeeper water—these just above a non-migratory populationof carnivorous Cyclothone. The carnivorous medusa Solmissusalbescens moved throughout the upper 600 m in the course ofits diel vertical migration. Offshore, carnivores were dominantthroughout the water column, with numerous diphyid siphonophoresin the upper layers, and Cyclothone, lobate ctenophores andmacroprotozoa abundant in deeper water. Solmissus was also present,and was more numerous offshore than in the coastal zone.     

The fate of discarded appendicularian houses: degradation by the copepod, Microsetella norvegica, and other agents

Despite the potential importance of zooplankton in degradationof marine snow, the association of colonising zooplankton withdiscarded appendicularian houses has not been investigated innorthern areas. We sampled the vertical distributions of appendicularians,houses and potential zooplankton colonisers at two stationsin the central North Sea during late summer. In addition, grazingexperiments were performed with the copepod Microsetella norvegica,which was assumed to be the main contributor to house degradation.The results were used in (i) inverse modelling, to estimatethe factors which were significant in shaping the vertical distributionof houses and (ii) calculations to estimate potential housedegradation rates. M. Norvegica was able to feed on appendicularianhouses, with feeding rates up to 0.42 g C (g C)^–1 day^–1(0.14 µg C ind.^–1 day^–1). The model resultssuggested that the vertical distribution of houses was shapedby sinking of houses, bacterial degradation and feeding of M.norvegica and invertebrate larvae. The estimated community degradationrate by M. norvegica was low, whereas invertebrate larvae haddegradation rates close to bacterial degradation. We concludethat at the typical concentrations of M. norvegica in the NorthSea (10⁴ ind. m^–2), its role in marine snow degradationis likely to be small. Degradation by other zooplankton groups,such as invertebrate larvae, can, however, be substantial.     

Spatial and temporal distribution of mesozooplankton in the Gulf of Aqaba and the northern Red Sea in February/March 1999

The distribution pattern, taxonomic composition and communitystructure of mesozooplankton was studied along a transect with10 positions between the Gulf of Aqaba and the northern RedSea. Five positions were resampled two or three times duringa cruise of RV ‘Meteor’ in February/March 1999.In spite of clear differences in the density stratificationbetween the Gulf of Aqaba and the northern Red Sea, the mesozooplanktoncomposition was very similar: Copepods were by far the mostabundant taxon, contributing 76–95% to the total community.The remainder was composed largely of ostracods, chaetognaths,appendicularians and molluscs. The mesozooplankton of the deeplymixed stations was homogeneously distributed, at all other stationsthe bulk of the mesozooplankton (>70%) was concentrated inthe mixed surface layer with peaks of calanoids, cyclopoidsand appendicularians in the vicinity of the chlorophyll a (Chla) maximum layer. Ostracods and poecilostomatoids dominatedthe layers below. Standing stocks within the total water column(550–1200 m) varied between 93 and 431 x 10³ individualsm^–2 for copepods and 5–76 x 10³ individuals m^–2for other mesozooplankton with highest numbers in the northernGulf of Aqaba, where vertical mixing was deep (400–500m) and Chl a and mesozooplankton distributions homogeneous throughoutthe water column. Towards the south, the mixed depth decreasedfrom 300 m in the central Gulf of Aqaba to 50 m in the Red Sea.Cluster analysis separated three distinct groups of stations,compounding the observed differences between the northern Gulfof Aqaba (Position I) and the other positions. The analysisalso revealed temporal differences between the February andMarch sections of the cruise, indicating the winter–springtransition. The stations sampled in March are characterisedby a higher total abundance and by a higher percentage of appendiculariansand ostracods than the stations sampled in February     

Productivity and grazing impact of Oikopleura dioica (Tunicata, Appendicularia) in Tokyo Bay

To estimate the productivity and grazing impact of a commonappendicularian species Oikopleura dioica in Tokyo Bay, monthlyobservations on its abundance and vertical distribution wereconducted during 2000. The abundance peaked in February andOctober, but was low during summer. Seasonal fluctuations inproductivity were similar to those of the abundance, with maximumvalues of 92, 134 and 508 mg C m^–2 day^–1 for somatic,new house and discarded house productivity in October and 206mg C m^–2 day^–1 for fecal pellet productivity inFebruary, respectively. The averaged biomass of O. dioica wasonly 2.5% of that of copepods; however, the secondary productivityof the former corresponded to 12.4% of the latter. Daily grazingimpact on particulate organic carbon ranged from 0.05% to 5%,which is close to the impact by copepods. These results implythat in Tokyo Bay, where small copepods and jellyfish are abundant,O. dioica is an important component of the ecosystem becauseit bridges between small primary producers and higher consumers.     

Effects of temperature and body size on the clearance rate of Oikopleura dioica

The effects of temperature and body size on the clearance rateof the appendicularian Oikopleura dioica were investigated.The clearance rate increased with the trunk length of the animalsaccording to a power function with exponent 3.2 ± 0.17(range 2.9–3.8). The intercepts of the three regressionswere significantly different with the highest rates at 20°C,corresponding to a Q₁₀ value of 1.78 (10–20°C).     

Repartition des Appendiculaires au large des cotes de Californie

Twenty-seven taxa of appendicularians have been identified fromthe cruise CALCOFI n° 7202 and their distribution has beenstudied by various numerical methods, with the purpose of definingsimultaneously the groups of associated species and the groupsof stations possessing similar characteristics. Two large recurrentgroups have been defined by the method of Fager and McGowan,corresponding approximatively to the major water masses present:central Pacific waters and equatorial Pacific waters. The methodof Williams and Lambert permits the separation of 11 southernstations and a northern group divided into 2 subgroups dependingon the presence or absence of O. dioica. The rank correlationsgive results very similar to those obtained with point correlations.Finally, principal component analysis allow good separationof the northern and southern zones by the factorial plan 1–3,while representation of the stations by the factorial plan 2–3separates the two northern subgroups depending on the presenceor absence of O. dioica. The results obtained by the differentmethods are therefore very much alike, but it is difficult toconclude they are a result of the specific cruise, or whetherthey represent a general phenomenon.     

中国贝母属(Fritillaria)7种植物的核型研究

通过对7种国产贝母属（Fritillaria）植物的染色体核型进行观察和研究,报道了小白花贝母（F.albidoflora Duan&Zheng）、川贝母（F.cirrhosa Don）、伊贝母（F.pallidiflora Schrenk ex Fischer&Meyer）、华西贝母（F.sichuanica Chen）、托里贝母（F.tortifolia Duan&Zheng）、新疆贝母（F.walujewii Regel）、裕民贝母（F.yuminensis Duan）等7种植物的染色体数目及核型,其中3种为首次报道。结果显示,7种国产贝母属植物的核型均具有高度不对称性。此外,小白花贝母与已报道的黄花贝母（F.verticillata Willdenow）的核型存在明显差异,提示Flora of China将小白花贝母归并入黄花贝母的分类处理可能并不恰当,二者的关系需进一步研究。     

Photosynthetic Characteristics of C3-C4 Intermediate Flaveria Species II. Kinetic Properties of Phosphoenolpyruvate Carboxylase from C3, C4 and C3-C4 Intermediate Species

The kinetic properties of phosphoenolpyruvate (PEP) carboxylasehave been studied among several Flaveria species: the C₃ speciesF. cronquistii, the C₃–C₄ species F. pubescens and F.linearis, and the C₄ species F. trinervia. At either pH 7 or8, the maximum activities (in µmol.mg Chl^–1.h^–1)for F. pubescens and linearis (187–513) were intermediateto those of the C₃ species (12–19) and the C₄ species(2,182–2,627). The response curves of velocity versusPEP concentration were hyperbolic for the C₃ and C₃–C₄species at either pH 7 or 8 while they were sigmoidal for theC₄ species at pH 7 and hyperbolic at pH 8. The K_m values forPEP determined from reciprocal plots were lowest in the C₃ species,and of intermediate value in the C₃–C₄ species comparedto the K' values of the C₄ species determined from Hill plotsat either pH 7 or 8. Glucose-6-phosphate (G6P) decreased theK_m values for PEP at both pH 7 and 8 in the C₃ and C₃–C₄species. In the C₄ species, G6P decreased the K' values at pH8 but increased the K' values at pH 7. In all cases, G6P hadits effect by influencing the activity at limiting PEP concentrationswith little or no effect on the maximum activity. At pH 8 andlimiting concentrations of PEP the degree of stimulation ofthe activity by G6P was greatest in the C₄ species, intermediatein F. linearis, a C₃–C₄ species, and lowest in the C₃species. In several respects, the PEP carboxylases of the C₃–C₄Flaveria species have properties intermediate to those of theC₃ and C₄ species. (Received April 30, 1983; Accepted August 22, 1983)