Behavioral interactions between a cyclopoid copepod predator and its prey

Behavioral observations on the predatory interactions betweenMesocyclops edax and several different types and sizes of preyrevealed that prey size alone was less important than otherspecific morphological and behavioral characteristics of theprey in deterring successful predation by the copepod. The behavioralresponses of Bosmina and Asplanchna to an attacking copepodwere passive and consisted of a simple retraction of vulnerableswimming appendages which made the prey more difficult to grasp.Daphnia and Diaphanosoma on the other hand exhibited very activeswimming escape responses. Tropocyclops usually avoided M. edaxby fleeing before the larger predator could detect them. Thehard carapaces of Daphnia, Bosmina and Keratella were effectiveat reducing ingestion following capture by M. edax. The resultsof these behavioral observations were supported by enclosureexperiments in which the predator was offered a choice betweentwo prey simultaneously. Cyclopoid copepods are capable of successfullyattacking, capturing and ingesting prey organisms several timestheir own body length. Although size alone may influence thepreference of cyclopoid copepods on large and small individualsof the same or similar prey species, it is not a dependabledeterminant of the preference of cyclopoids on multispecificprey assemblages. ¹Present address: Department of Biology, Williams Hall No. 31,Lehigh University, Bethlehem, PA 18015, USA     

Diel periodicity and selectivity in the feeding rate of the predatory copepod Mesocyclops edax

The diel periodicity and selectivity in the feeding behaviorof the predatory cyclopoid copepod Mesocyclops edax was investigatedat 3 h intervals over two 24 h sampling periods in nature. Gutcontent analyses revealed an increase in gut fullness at 11.30and 20.30 in July, and at 20.30 in August. The increase in gutfullness at these times could not be explained by an increasein prey density, changes in predator-prey overlap or differencesin prey vulnerability. We suggest that M.edax exhibits a truediel periodicity in the intensity of its predatory activities,although the alternative hypothesis of a diel periodicity ingut passage time cannot be ruled out. Vanderploeg and Scavia'sE^* selectivity index indicated a preference for rotifer andcladoceran prey over copepods.     

Dynamics of a subtropical plankton community,with emphasis on the copepod Mesocyclops edax

The zooplankton community of a subtropical, eutrophic lake was studied over an annual cycle. Mesocyclops edax, the numerically dominant crustacean, exhibited a stationary age distribution for most of the year (July–March). Per capita growth rates (r) of M. edax approximated zero throughout most of the study, indicating a close correspondence between birth and death rates. High death rates in the spring and summer probably were due to intense predation, since physiological mortality was low and predators were abundant. Multiple regression analyses indicated that much of the variation in M. edax birth rates could be explained by abundance of their animal prey. Other plankton in this lake exhibited considerable seasonality, but not to the marked degree characteristic of temperate populations. The number of zooplankton species was low and cladocerans were scarce, as is characteristic of many eutrophic Florida Lakes.     

Environmental factors influencing diurnal distribution of zooplankton and ichthyoplankton

The diurnal vertical distribution of a large number of speciesof zooplankton, icbthyoplankton and micronekton were determinedin the top 150 m in three locations in the Shelf Water, on theNova Scotia Shelf, and Slope and on Georges Bank during springand fall periods. Species were categorized as to their trophiclevel and their type of diurnal migration behaviour. The influenceof temperature, salinity, and water density on the diurnal verticaldistribution of the species was examined. Temperature was foundto have the greatest influence on the distribution of the largestnumber of species. Diurnal migration behavior of the same speciesin Shelf and Slope water and at different times of the yearwas examined. Results showed that species changed their behaviorin the two water masses, while some species changed their migrationbehavior at different times of the year. During the night inApril the most abundant copepod species, Calanus finmarchicus,making up about 80% of the biomass, was found concentrated abovethe thermocline and the main chlorophyll layer. The majorityof the less abundant species of copepods were found below thethermocline and the chlorophyll layer. At night in August thetwo most abundant copepod species, Centropoger typicus and Paracalanusparvus, making up at least 80% of the zooplankton biomass, werealso concentrated above the thermocline and the main chlorophyllLayer. Three species of copepods were concentrated at the depthof the main chlorophyll layer and two species were concentratedbelow the chlorophyll layer and thermocline. The vertical distributionof other zooplankton and ichthyoplankton species was examinedin relation to the thermocline and chlorophyll layer. Relationshipsbetween concentrations of six species of fish larvae and allspecies of copepods in the same samples showed a general increasein the numbers of larvae m^–3 as the numbers of copepodsm^–3 increased in a range of 500–4000 m^–3.However, the concentration of Merluccius bilinearis decreasedas the concentration of copepods exceeded 4000 m^–3 suggestingthat high concentrations of copepods may not be a favourableenvironment for the larvae.     

Morphological responses by Bosmina longirostris and Eubosmina tubicen to changes in copepod predator populations during a whole-lake acidification experiment

Changes in zooplankton populations during the experimental acidilicationof Little Rock Lake provided an opportunity to examine specificmechanisms underlying the morphological responses of bosminidsto changing predation pressure. Two large copepods, Epischuralacustris and Mesocyclops edax, disappeared from the lake'sacidified basin in 1986 and 1989, respectively, while a smallercopepod predator, Tropocyclops extensus, increased during laterstages of acidification. The two bosminid species showed distinctlydifferent responses coinciding with the changes in copepod predation.Bosmina longirostris exhibited a significant decrease in mucrolength with the decline of M.edax and E.lacustris. Its meanbody and antennule length, however, did not change. We suggestthat the decoupling of B.longirostris mucro length and antennulelength may have been related to the persistence of the smallercopepod predator, T.extensus. Eubosmina tubicen showed no apparentresponse to declines in M.edax and E.lacustris abundance ineither mean mucro, antennule or body length. Allometric analysesindicated, however, that mucro length was related to size-dependentcopepod predation for both B.longirostris and E.tubicen.     

The nutritional ecology of the ctenophore Bolinopsis vitrea: comparisons with Mnemiopsis mccradyi from the same region

Bolinopsis vitrea is a warm water lobate ctenophore which doesnot overlap in its distribution with Mnemiopsis mccradyi incontiguous waters. We examined its feeding ecology on a seriesof cruises. B. virrea ingested increasingly more prey at higherfood concentrations (2–100 prey l^–1) but feedingeffort (clearance rate) decreased with increasing food availability.On a dry weight basis, smaller tentaculate Bolinopsis ingestedseveral times more than larger lobates, but based on carbonweight, specific ingestion was fairly uniform over the entiresize range investigated (6–60 mm total length). Bolinopsiscollected during the daytime in the Bahamas rarely had morethan three prey items in their guts. These results and laboratorymeasurements of digestion times (av. = 1.9 h) allowed computationof daily rations, which could not account for the metabolicrequirement as measured on the same cruises. Results of feedingexperiments, however, implied that prey densities in excessof 11^–1 were sufficient to sustain a growing populationof Bolinopsis. Prey concentrations about an order of magnitudehigher were required for M. mccradyi based on similar experiments.These results were in general agreement with observed densitiesand distributions of ctenophores and their zooplankton preyin the Bahamas and coastal South Florida.     

Diel periodicity of Subantarctic copepods: relationships between vertical migration, gut fullness and gut evacuation rate

A zooplankton community in the Polar Frontal Zone north of SouthGeorgia was sampled for 5 days in February 1994. Feeding ofvarious copepodite stages of six copepod species was assessedby a series of gut fluorescence/gut evacuation experiments.Feeding periodicity was compared to vertical distribution andmigration patterns, as revealed by Longhurst Hardy PlanktonRecorder (LHPR) and ring net catches. Despite chlorophyll alevels reaching only 0.8 mg m^–3 and daily carbon rationsbased on phytoplankton intake being low, feeding of all thecopepods was restricted mainly to the 8 h night-time perid.During the daytime, the epipelagic community was verticallydispersed within the top 100 m. At night, upward migration bymost species led to a convergence of almost all zooplanktersin the upper half of the surface mixed layer. However, large-amplitudevertical migrations which crossed the thermocline were performedonly by Metridia lucens and Pleuromamma robusta. Although feedingby both migrants and non-migrants was mainly at night, therewas no diel signal in gut evacuation rate. The gut evacuationrates of the nine copepod species and stages differed significantly(5-fold) and were negatively related to the extent of theirdiel vertical migration. The long gut-passage times of the migratoryspecies, M.lucens and P.robusta, would have allowed them timeto defaecate some of the food eaten near the surface at depth,contributing to an active carbon transfer out of the mixed layer.However, their scarcity at this site meant that their grazingcomprised only -<1.4% of total copepod ingestion, so theircombined gut flux is likely to have been negligible (-0.4 mgcarbon day^–1).     

Omnivory in cyclopoid copepods: comparisons of algae and invertebrates as food for three, differenfly sized species

We examined the importance of algal versus invertebrate preyfor three cyclopoid copepods by comparing reproductive successfor females fed on three diets: exclusively algal food, exclusivelyinvertebrate prey and a combination of algal plus invertebrateprey. The three cyclopoid species represent a distinct gradientin body size: Tropocyclops prasinus mexicanus (small body size),Diacyclops thomasi (medium body size) and Mesocyclops edax (largebody size). Our results provide strong evidence for the potentialimportance of algae in the diet of adult cyclopoid copepods.However, the role of herbivory is dependent on species bodysize. We found that algae are the key dietary factor for reproductivesuccess of the small T.p.mexicanus. In contrast, the availabilityof invertebrate prey had a greater influence on reproductionby the two larger species. Overall, cyclopoid copepods shouldbe generally considered as having the potential to derive theirenergy from either animal or algal sources. This has importantconsequences for interpretations of the forces influencing zooplanktoncommunity diversity as well as for general food web theory.     

Photoresponses of the copepod Mesocyclops edax

The predatory copepod Mesocyclops edax is an important componentof many zooplankton communities where it typically makes extensivedid vertical migrations. To describe the effect of light onadults we measured their photoresponses in the laboratory. Theresponse spectrum is characterized by a wide plateau of greatestsensitivity from about 480 – 580 nm. These animals areadapted to perceive light during the day since their regionof maximum sensitivity overlaps the spectral region of highestquantal intensity underwater (575 – 700 nm). The thresholdintensity for positive phototaxis by dark adapted animals wasabout 5 x 10^–1 Wm^–2 at 540 nm, and they were positivelyphototactic up to an intensity of 5 x 10^–1 Wm^–2.Above this intensity phototaxis is no longer observed. Light-adaptedanimals were less sensitive than dark-adapted, but their generalpattern of response to light intensity did not differ. Thereis no rhythm in phototaxis. Their photoresponses may providea mechanism for controlling vertical migration so as to minimizeexposure to planktivorous fish. ¹Contribution No. 1375-AEL from UM-CEES, Appalachian EnvironmentalLaboratory.     

Interannual changes in the biomass of the Black Sea gelatinous zooplankton

The 15 year changes in the total gelatinous biomass consistof a general trend to increase, from 250 gm^–2 in 1980to 2500 g m^–2 in 1995, and fluctuations with a periodof –4 years performed against the background of this trendin the upper 150 m layer. Different species occupying distincttrophic niches form these peaks. When represented as the percentageof the total zooplankton biomass, Aurelia aurita and Mnemiopsisleidyi exhibit the contra-phase fluctuations where a sharp increasein M.leidyi is accompanied by a decrease in the A.aurita biornass.     

Planktivory by larval Odontesthes bonariensis (Atherinidae: Pisces) and its effects on zooplankton community structure

This paper describes the effect of predation by larvae of theatherinid Odontesthes bonariensis upon the zooplankton communityduring a 3-month experiment Three-day-old larvae were stockedin 45 m² concrete tanks at rates of 100 and 200 fish m^–2Gut content analyses showed that the larvae consumed relativelysmall prey all along the experiment Morphological (mouth width)and physiological (gastric inefficiency) constraints seems tohave precluded the capture of the largest prey. The prey community,however, showed all the symptoms generally ascribed to size-selectivepredation upon the largest individuals: decrease in maximumzooplankton size and mean cladoceran size, decrease in cladoceranand copepod biomass, extinction of the largest zooplankton species(Daphnia stmilis), increase in rotifer biomass, etc It is concludedthat size-selective predation is not a necessary condition forthe commonly observed decrease in zooplankton size after theincrease in density of a vertebrate predator. The ecologicalimplications of this result are discussed. Present address: Department of Biology, Lehigh University, WilliamsHall 31, Bethlehem, PA 18015-3189, USA     

DISTRIBUTION AND ABUNDANCE OF GYMNOSOMATA (GASTROPODA: OPISTHOBRANCHIA) IN THE SOUTHWEST ATLANTIC

The distribution and abundance of gymnosome gastropods in theArgentine Sea and Brazil—Malvinas Confluence during 1978–1979and 1988 were studied. The collections analyzed included 768quantitative samples obtained between 48°W and the coast,and from 35°S to 55°S. Two species were found. Spongiobranchaeaaustralis was the most frequent and abundant (up to 730 per1000 m3); its presence in the area was associated with the coreof the Malvinas Current. Clione antarctica was less abundant(maximum abundance: 230 per 1000 m³) and was also associatedwith the Malvinas Current. The geographic ranges of both speciesin the area are wider than previously described. Since the rangeof S. australis in the area extends far from the range of itsprey Clio, it is not clear whether S. australis can feed onthe thecosomatous pteropod Limacina (and not only on Clio, asdescribed in the bibliography) or it starves in that area. Duringthe 1978–1979 annual cycle, the abundance of both speciesfollowed neither the abundance patterns of their prey nor ofthe total zooplankton, and differed from each other. The residencetime of swarms of both gymnosomes were shorter than one month.As a general pattern, the aggregates are rapidly transportednorthward by the Malvinas Current and also penetrate the outershelf water, but they remain there only during a short periodand cannot preclude the final expatriation. So, the abundanceof gymnosomes in the area depends on passive migration morethan intrinsic population factors. (Received 9 July 1997; accepted 15 December 1997)     

Structure and dynamics of zooplankton communities maintained in closed systems, with special reference to the algal food supply

Seasonal changes in the structure of the zooplankton communitiesof two large (18 500 m³) experimental enclosures (‘LundTubes’) are reviewed. The diets of the two most prominentcrustaceans, Daphnia hyalina and Diaptomus gracilis, were investigatedby direct analyses of gut contents and in a series of shortterm experimental manipulations of communities isolated in 6m³ bags, placed in one of the enclosures. The relative abundanceof Daphnia hyalina generally coincided with warmer water temperaturesbetween late spring and late summer, but its numbers fluctuatedwithin that period, apparently in response to the availabilityof smaller algae (<10⁴ µm³ unit volume: chiefly cryptomonads,chlorellids and Ankyra). Daphnia numbers declined steadily throughthe summer in the enclosure overwhelmingly dominated by Microcystis,although it did feed on smaller colonies (< 10⁵ µm³in volume). Diaptomus gracilis and most rotifers were also prominentin the first half of the year, but peak abundances of Keratellaquadrata followed the presence of large populations of small(<10³ µm³) algae. Under conditions of limiting foodavailability, mortality was greatest amongst the smaller speciesand the juvenile instars of the large Crustacea. ¹Present address: Anglian Water Authority, North Street, Oundle,Peterborough, UK     

Vertical distribution, life cycle and production of the chaetognath Sagitta crassa in Tokyo Bay, Japan

The daytime vertical distribution of Sagiita crassa in TokyoBay was examined from February 13, 1988 to February 20, 1989.High densities of larger-size chaetognaths were found near thesea bottom, whereas the smaller animals tended to inhabit theupper layers. This feature of distribution is discussed in relationto the distribution of their main food organisms, e g. Pseudodiaptomusmarinus, Acartia omoru, Centropages abdomialis and Oithona davisae.The two periods of replacement of two morphs were confirmedby the variation only in mean body length of this chaetognath,unlike the previous authors who made additional morphologicalobservations. It was hypothesized that S.crassa has at leastfive generations Two generations, including mostly the largerforms, had higher growth rates than the generation consistingmainly of the small form. Yearly respiration of S.crassa was8.2 g C m^–Abstract. Yearly production of this animal wasestimated to be 3.8 g C m^–. A feeding estimate revealedthat chaetognaths require a prey production of 13.1 g C m^–year^–1. The impact of this chaetognath on the prey populationin Tokyo Bay and the propriety of an estimated value of annualproduction of S crassa is discussed.     

Biogeographic study of the planktonic communities of the Prince Edward Islands (Southern Ocean)

Microphytoplankton and zooplankton composition and distributionin the vicinity of the Prince Edward Islands and at the Sub-antarcticFront (SAF) were investigated in late austral summer (April/May)1996. Samples were collected for analysis of chlorophyll a concentration(Chi a), microphytoplankton and zooplankton abundance. Generally,the highest Chl a concentrations (up to 2.0 µg l^–1)and zooplankton densities (up to 192 ind. m^–3) were recordedat stations within the inter-island area while the lowest values(<0.4 µg l^–1) were observed at stations upstreamof the islands. High Chl a and zooplankton biomass values werealso associated with the SAF. Microphytoplankton were dominatedby chain-forming species of the genera Chaetoceros (mainly C.neglectus),Fragilariopsis spp. and the large diatom Dactyliosolen antarcticus.The zooplankton assemblages were always dominated by mesozooplanktonwhich at times contributed up to 98% of total zooplankton abundanceand up to 95% of total biomass. Among mesozooplankton, copepods,mainly Clausocalanus brevipes and Metridia lucens numericallydominated. Among the macrozooplankton euphausiids, mainly Euphausiavallentini, E.longirostis and Stylocheiron maximum, and chaetognaths(Sagitta gazellae) accounted for the bulk of abundance and biomass.Cluster and ordination analysis did not identify any distinctbiogeographic regions among either the microphytoplankton orzooplankton.     

Experimental evaluation of herbivory in the ctenophore Mnemiopsis leidyi relevant to ctenophore-zooplankton-phytoplankton interactions in Narragansett Bay, Rhode Island, USA

Herbivory of Mnemiopsis leidyi and its interactions with phytoplanktonand non-gelatinous zooplankton were examined in small-scalemicrocosm experiments. Clearance rates for M. leidyi incubatedwith phytoplankton were generally negative, but ranged up to4.5 1 ctenophore^–1 day^–1 when the large (80 µmø) diatom Ditylum brightwelli was offered as a food source.These highest ingestion rates would provide Mnemiopsis withonly 21 % of its daily carbon requirements for respiration.Mean shrinkage of M. leidyi was 8.2–51% when incubatedwith phytoplankton. Although M. leidyi neither fed activelyon phytoplankton, nor satisfied its nutritional needs on sucha diet, the chain-forming diatom Skeletonema costatum becameentangled in mucus strands and balls produced by M. leidyi inthe absence of zooplankton. Attachment onto mucus occurred atphytoplankton concentrations commonly observed in NarragansettBay and may be important in the formation of "marine snow" duringsummer M. leidyi pulses; phytoplankton sinking rate and the"package size" available to herbivores would also be affected.The experiments support our previous hypothesis based on fieldobservations in Narragansett Bay that M. leidyi indirectly regulatesphytoplankton abundance there during the summer bloom as a consequenceof predation on zooplankton. The extent to which M. leidyi influencedphytoplankton dynamics in the microcosms was dependent on therelative abundance and physiological state of the three trophiclevels. A food web diagram for M. leidyi is presented.     

Zooplankton prey selection by juvenile fish in Nova Scotian Shelf basins

The stomach contents of fish <100 mm standard length (SL)and fish >100 mm SL collected during June and October intwo deep basins on the Scotian Shelf, showed that juvenile co(and silver hake fed primarily on the two most abundant speciesof zooplankton in the basins: Calanus finmarchicus and Meganyctiphanesnorvegica. These zooplankton were abundant in the basin: duringthe 2 months, forming high concentrations at depths below 200m during the day. In June fish (<100 mm SL) concentratedin the upper 50 m during both day and night. Cod and wolffislcollected between 195 and 240 m had a large percentage of C.finmarchicusin their stomachs suggesting that these fish exploited the highconcentrations of C.finmarchicus below 200 m depth There wasno evidence that Calanus hyperboreus was eaten by any juvenilefish species. Calanus finmarchicus was the most common preyof cod feeding in the top 50 m, but the percentages o C.finmarchicusstages IV and V were lower in the stomachs than in the watercolumn. Cod showed ; strong preference for M.norvegica, withthis prey species having a modified Ivlev's index of 1 indicatingthat cod were highly selective for this species. There was noevidence that C.hyperboreus. stage IV was eaten by any of thespecies offish. Silver hake and dogfish (>100 mm SL) bothfeed oi M.norvegica and C.finmarchicus in the deep regions ofthe basins. In October, silver hake was the most common juvenilefish in the basins, feeding primarily on small stages of M.norvegica.The populations of fish (>100 mm SL) in the basins were mainlyspecies known to feed heavily or euphausiids. The two basinsare unique regions of the shelf because of the large populationsof Calanus copepods and M.norvegica they contain all the yeararound, thereby providing an atrractive feeding ground for manyspecies of fish, particularly silver hake.     

Zooplankton, especially calanoid copepods, in the upper 1000 m of the south-east Arabian Sea

Abstract. Vertical distribution of zooplankton in the upper1000 m was studied from the south-east Arabian Sea in orderto determine the variations in zooplankton at different depths.The distribution and migration patterns of calanoid copepodspecies were given special attention. The mean zooplankton standingstock in the upper 1000 m was 2.1 g dry Wt m^–2, of which97.7% was concentrated in the upper 400 m. Herbivores were generallymore abundant at all depths, but did not predominate. An increasein zooplankton at night occurred in the upper 200 m, as wellas at the 600–1000 m stratum. The maximum diversity ofcalanoid copepods also coincided with these two depths. Basedon vertical ranges, the calanoid copepod species were assignedto three groups: (i) species occurring predominantly in theepipelagic layer and forming the bulk of the calanoids; (ii)relatively sparser deeper living species confined below 200m; and (iii) species occurring throughout the water column.Some amount of vertical niche separation among congeneric specieswas indicated. While some species showed active migration, agood number of species were non- migratory.     

The influence of food resources on the development, survival and reproduction of the two cyclopoid copepods: Cyclops vicinus and Mesocyclops leuckarti

Laboratory experiments were conducted to determine whether thetwo cyclopoid copepods. Cyclops vicious and Mesocyclops leuckaru.exploit the same food resources. The food requirements of juvenilesof the two cyclopoid copepods were investigated. Moreover, theimportance of algae for the predaceous adults was studied. Naupliiof both M leuckaru and C.vicinus successfully developed intocopepodites when fed the motile algae Chlamydomonas reinhardtii.Chlamydomonas sphaeroides and Cryptomonas sp. Threshold foodconcentrations for naupliar development varied between offeredalgae and between the two cyclopoid species. The food thresholdfor successful naupliar development, when reared on C.reinhardui,was lower for M.leuckarti (0.3 mg C 1^–1) than for C.vicinus(0.5 mg C l^–1) whereas a similar food threshold was foundusing Cryptornonas sp (0.3 mg C ^–1) and C.sphaeroides(<0.2 mg C 1^–1), Naupliar development time was inverselyrelated to food concentration. Food required for copepoditedevelopment differed for the two cyclopoid species. Cyclopsvicinus was able to develop to the adult stage on a pure dietof any one of the three algal species. whereas M.leuckarti requireda prey supply of the rotifer Brachionus rubens. Food composition.i.e. algal species, algal concentration and rotifer abundance,influenced copepodite survivorship of both cyclopoids and wasalways higher in the presence of B.rubens. Under similar foodconditions, mortality was higher for M.leuckarti than for Cvicinus. Mesocyclops leuckaru females were very dependent onanimal food. The predation rate of M.leuckaru was not lowerin the presence of algae. Egg production of M.leuckarti waslow on a pure algal diet and significantly higher when B rubenswas present. The results were used to discuss the life cyclestrategy and the possibility of exploitative competition ofthe two cyclopoid copepods.     

The impact of short-term exposure to UV-B radiation on zooplankton communities in north temperate lakes

We exposed natural zooplankton communities to in situ levelsof sunlight for 3 days at different depths in two north temperatelakes: one oligotrophic and one eutrophic. Natural fluxes ofUV-B radiation (280–320 nm) were manipulated with cutofffilters (Mylar®). There was substantial mortality in someof the zooplankton exposed to UV-B in the oligotrophic lake,but not in the eutrophic lake. Reproduction of Diaptomus wassuppressed by UV-B down to 6 m in the oligotrophic lake. Thesedata suggest that natural levels of UV-B radiation in the oligotrophiclake may prevent some species of zooplankton from continuouslyexploiting the warm surface waters during summer stratification.In the more eutrophic lake, UV-B is a less important constraintin the vertical distribution of zooplankton. These differencesin the responses of zooplankton to natural UV-B radiation inlakes may alter their ecological interactions with food resources,predators and other environmental variables in the water column. ¹Present address: Universidad Nacional del Comahue, Centro RegionalUniversitario Bariloche, CC 1336, (8400) Bariloche, Argentina ²Present address: National Academy of Engineering, 2101 ConstitutionAvenue, NW, Washington, DC 20418, USA