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MARINE PROGRESS SERIES Published September 25 Mar Ecol Prog Ser

The bioluminescent field of the Atlantic

S. A. ~iontkovski',Yu. N. Tokarevl, E. P. Bitukovl, R. ~illiarns~~*,D. A. ~iefer~

'Institute of of the Southern , 335011 Sevastopol, Ukraine 2~lymouthMarine , Prospect Place, Plymouth PL1 3DH. United 3Department of Biological Sciences, University of Southern California. Los Angeles, California 90089-0371, USA

ABSTRACT: Data from 20 yr (1970 to 1990) of expeditions by the Institute of Biology of the Southern Seas, Ukraine, to the tropical are summarised in the form of a macroscale contour map. The bioluminescent intensity of in the upper 100 m layer was analysed from 2924 casts. Sev- eral zones of enhanced bloluminescence are shown from the annual averages (0 to 100 m),associated with major along the African and geostrophic currents formlng the westward mass transport. The macroscale trend of spatial distribution and the stochastic component of the biolu- minescence were partitioned by analysis of the autocorrelation functions. General agreement between and biornass distributions was noted on an ocean basin scale. The con- tribution of phyto- and zooplankton fractions to the formation of the integrated bioluminescence poten- tial can vary significantly in the upper 100 m layer over regions within the tropical zone.

KEYWORDS: Plankton biolumincscence . Spatial heterogeneity Atlantic Ocean

INTRODUCTION The bioluminescence intensity or potential, mea- sured for the whole , can also act as an indi- The integrative structural and functional biological cator of plankton , and functional characteristics of the oceanic biota, like the phyto- and state of planktonic (Marra & Hartwig 1984, zooplankton biomass, a concentration and Losee et al. 1989, Marra 1989, Evstigneev & Bitukov , are widely used in multipurpose 1990, Lapota et al. 1994). The plankton biolumines- studies of marine ecosystems. The numerous long- cence, however, is among those integrative character- term measurements of these characteristics in various istics which have never been amalgamated or repre- geographical regions have enabled macroscale gener- sented in a form of the map, based on direct field alisations to be made and maps of planktonic fields of measurements, for any large-scale region or the ocean. the world ocean to be created (Bogorov et al. 1968, There are a number of reasons for this situation. Firstly, Koblentz-Mishke 1977, Semina 1977). Such maps have the majority of ongoing studies of bioluminescence in been progressively revised during recent decades for a the were, and still are, located in the coastal zones. number of regions (Chromov 1986, Piontkovski & Secondly, it was mainly the former Soviet Union whose Ignatyev 1992, Rudjakov & Tseitlin in press). Together projects were associated with the long-term surveys of with data on hydro-optical characteristics and water the open of the world ocean (1970 to 1990). mass dynamics, such generalised observations serve as However, these data were restricted in their publica- the basis for biogeographical partitioning of the tion and were generally inaccessible. The international (McGowan 1971, Van der Spoel & Heyman 1985, cooperation programmes of data exchange between Longhurst et al. 1995). the former Soviet Union, and the USA (IOC 1992) have enabled data to be amalgamated and jointly analysed. 'Addressee for correspondence. We present the map of the Atlantic Ocean biolumi- E-mail: [email protected] nescence field for the first time.

0 Inter-Research 1997 Resale of fullarticle not permitted Mar Ecol Prog Ser 156 33-41, 1997

Submersible gear. The biolumlnescence intenslty vds measured by means of the bathyphotometer 5 4LP\ (Vasilenko et a1 1997) with a dynamic band of 57 decibars The bioluminescence sensor was mounted at the bottom of a cylinder and detected the bloluml- nescence emltted b.i. wlth~na dark chamber This dark chamber had a protect~ngscreen and 4 impellers rotatlng on a fluoroplastlc bea~ingThe pitch of the lmpeller vanes protected the chamber from ex- ternal llght When the Instrument IS lowered the im- pellers are rotated by the flow of water The rotating lmpeller vanes mechanically dlsturb the plankton or- ganisms passlng through the dark chamber, causlng them to b~olum~nescepriol to measurement by the blo- lurnlnescence sensor The electrical slgnal from the sensor is transmitted up a single cable wlth temporal partitlonlng of the channels, to the deck unit The sensi- tlvity of the b~olumlnescencesensor is in a range from 10 l3 to 10 W cm wlth a band of spectral sens~t~vity from 160 to 600 nm The repeat tlme of the blolurnl- nescence temperature and conductivity sensors was 180 ms The bathyphotometer was deployed vertically at a speed of 1 3 m S-' The b~oluminescenceof plank- tonic organisms has a total latent perlod and flash dura- tion less than 0 6 S (Evstigneev & Bitukov 1990) whlch means that the sensor can easlly measure the blolumi- nescence s~gnalThe biolumlnescence signals coming from the submersible unlt were recorded and averaged

(0 'O l0 O l0 !O over 5 S intervals To calibrate the b~oluminescence sensor a radloactlve-luminescent llght source was used Fig 1 Posltion of statlons with vert~calcasts tor b~olum~nes cence and the 4 stations given In Table 2 The light intenslty of th~sstandard source was 12 X 10.' W A measure of the background llght was taken as the m~nimalblolumlnescence slgnal from 1 m depth METHODS The depth of the submersible gear was estimated by means of the pressure sensor DDV (precision 0 01 MPa) The b~oluminescencepotentlal or bloluminescence in- tenslty was the major parameter in the long-term fleld Table 1 General charactenstics of data used for the mapping measurements in the ocean This potential 1s the maxi- of blol um~nescence mum amount of radlant emitted In a glven vol- ume of water by stimulated bioluminescent organisms Month Number Number This can be estimated as B, = f B(t)dt where B(t)is the of stat~ons of prof~les light lntenslty during the biolumlnescent flash (At) Data were obtained from 15 expeditions of the Ukraln "13-Sep 12 lan Academy of Sciences to the Atlantic Ocean (Fig 1, Mar-Jun 26 Dec Var 2 1 Table 1) Field surveys consisted of drift statlons with Feb >Jar 11 measurements of biolumlnescence potentlal down to Dcc Var 42 200 m depth temperature, salin~ty,density profiles to Apr-Jul 57 1000 m depth, and vertical hauls by Juday plankton nets Dec-Jan 4 1 (36 and 82 cm mouth diameter, 142 to 220 pm mesh) Jul- Aug 4 1 Dcc-Feb 3 8 stratlf~edor integrated over 100 m depth Jan-Apr 442 Field measurements of biolumlnescence (2924 vertl- Dec-Jdn 553 cal casts conducted at 523 stations) were mainly con- Jul-Oct 793 ducted wlthln a penod from 2 h after sunset to 2 h Nov- Jan 626 Jun-bep before sunnse, to exclude the Impact of the vertlcal 66 Oct-Dec 155 migrations of the plankton and the llght die1 rhythm Piontkovski et al.. The bioluminescent field of the Atlantic Ocean 35

The absolute errors of the other sensors were O.Ol°C for temperature and 8 X 10-4 cm m-' for the conductivity channel. The integration time of the bioluminescence sensor was manually regulated in a band from 0.5 to 5 S. An 'ISTOK' CTD (with a temperature sensitivity of 0.0025"C, a measurements error of O.OOl°C, and a time lag for the sensor of about 50 ms) was used for vertical profiling. The specific electrical conductivity was 1.3 to 7.0 cm m-', the sensitivity was 0.0025 cm m-' with a measurement error of 0.002~(where X is the conductiv- ity number). The hydrostatic pressure was estimated by means of the pressure sensor with a band of 0 to 60 MPa (0 to 6000 dbar), a sensitivity of 0.004 to 0.007 MPa and a measurement error of 0.25 %. Data processing. To map the biolurninescence poten- tial of the upper 100 m layer the bioluminescence num- bers in the vertical profiles were averaged. The Surferm Golden Software Inc. (USA) package was applied to design the macroscale field of the biolun~inescence potential. The Kriging gridding method (Cressie 1991) and smoothing of the grid matrix, with the weight of matrix center = 2 and the distance weighting power = 2, were applied to create the bioluminescence map. Two- dimensional spatial autocorrelation functions were used to estimate the statistical structure of the stochas- tic (subgrid) component of the bioluminescent field. Phyto- and zooplankton fractions of plankton sam- ples were treated to the level of . The abun- dance of the bioluminescent, the probably biolumines- cent and non-bioluminescent species of both fractions were defined on the basis of a bioluminescent species list summarised in a number of reviews (Clarke et al. 1962, Lapota & Losee 1984, Evstigneev & Bitukov 1990, Gitelzon et al. 1992, Evstigneev et al. 1993).The prob- ably bioluminescent species are those species whose bioluminescence is still a subject of debate. To estimate the degree of similarity in changes of the bioluminescence potential, principal component ana- lysis was applied to the phyto- and zooplankton abun- dance. Parameter numbers were subjected to log(x+ 1) transformation. The rotated loadings and the percent- age of variance explained by components were de- fined on the basis of the VARIMAX procedure. The Pearson correlation matrix and linear regression analy- sis were used to estimate statistical links between bio- Fig. 2. Typical vertical profiles of bioluminescence intensity luminescence potential and above-defined log(x + 1) and temperature from vanous regions. (a) 1: 0" 30' S, l" 30' E transformed plankton components. (March 1981), 2. 28" 00' N, 39" 29' W (September 1982); (b)13" 19' N, 51" 43' W (February, 1974); (c)14" 30' N, l?" 55' W (March 1974).T: temperature (dashed lines); B,: biolumi- nescence intensity (10-\W cm-2 I-' ) RESULTS

The selection of the 100 m layer for processing and epipelagia lies in the upper 100 m layer. Patterns of averaging of vertical profiles was based on the prelim- vertical profiles (the number of bioluminescent peaks, inary analysis of the whole data set. This has shown their amplitudes, their position by depth, etc.) might be that about 90% of the bioluminescence potential of the distinctive for specific regions (Fig. 2). The compar- 36 Mar Ecol Prog Ser 156: 33-41, 1997

Fig. 4. Two-dimensional spatial autocorrelation function of the stochastic (subgrid) component of the bioluminescent field

Fig 3. Map of b~oluminescenceintensity (10-5 pW cm-' 1.') ot the Atlantic Ocean (0 to 100 m) The bioluminescent field of the tropical and subtrop- ical Atlantic Ocean exhibits some well developed and isons of the profiles of bioluminescence intensity with visually defined patterns. Firstly, there is a general those of phyto- and zooplankton a.llow us to conclude trend of decreasing bioluminescence intensity from the that the wide-bottom peaks of bioluminescence eastern to the western regions, especially from the (Fig. 2a) were mainly formed by increased abundance African regions towards South America. On of plankton organisms while the fine resolved peaks average, bioluminescence decreases 5 times or more are due to single flashes from or micro from 250 to 50-20 X 10-\W cm-' I-' (Fig. 3). swarms. In some cases profiles have (1)1 wide and The other feature is the less scaled va.ri.ations against well-defined maximum of bioluminescence usually the background of this general macroscale trend. The associated with the layer, (2) profiles have well-developed zones of enhanced bioluminescence several maxima or (3) the vertical profiles reflect the tdke the form of tongues streching west from the stratified biotope, where numerous layers of minimal African upwelling zones towards the open ocean. The and maximal bioluminescence are separated by sev- northern and central tongues cover regions of the eral metres. Finally, vertical statification of biolumines- Mauritanian coastal upwelling and the so-called cence might not be well developed and profiles might Guinea Dome. In the latter case, a quasi-stationary cy- have 1 or 2 low-amplitude peaks separated by 20 to clonic circulation takes place and could be monitored 30 m depth. The variety of vertical structure will cer- in a region with approximate coordinates 11" to 13" N, tainly affect the macroscale mapping of the bioluml- 21" to 22" W (Artamonov et al. 1987) The 'southern nescence at any of the horizons within the euphotic tongue' is more elongated along the African shelf. It layer but the maps we give integrate through all this covers the Benguela upwelling region and the variability. Analysis of the fine structure and diversity Angolan Dome, extends to the north and then curls of the vertical profiles is not within the scope of this westward near the equator. The central and western paper. regions of the South Atlantic have low productive oligo- Piontkovski et al.: The bioluminescent field of the Atlantic Ocean 3 7

trophic waters where bioluminescence 60 declines below 50 X lO-=pW cm-']-'. To evaluate the features of the sto- chastic component of the biolumines- cent field, the macroscale spatial trend 4,, and the field of the subgrid stochastic fluctuations were separated and ana- lysed on a statistical basis (Fig. 4). The 2-dimensional spatial autocorrelation function (ACF) of the pulsation or 20 stochastic component (characterising the synoptic scale processes) exhibits spatial anisotropy. Obviously biolumi- nescence fluctuations are statistically 0 linked on a space scale of about 5" to 7' along latitude and longitude. On the other hand, if we consider the so-called correlation radii of the ACF up to a 0.1 -20 correlation value, zones of correlated fluctuations of bioluminescence obvi- ously appear to be distributed along the equator and latitudinally in the direc- tion of the main water mass transport. -40 Spatial heterogeneity of the biolumines- cent field is also related to such synoptic events as frontal zones and current meanders. For example, in the South -60 -10 -20 0 2 0 Atlantic, contours of the 100 pW cm-' 1-' isO1ine the westward and Fig. 5. Spatial distribution of zooplankton biornass (0 to 100 m layer, mg m-3, the position of the macroscale divergent wet weight).Dots are sampling sites zones (see schemes of currents in Marti- nenko 1990). It is known that bioluminescence intensity can be and processed in detail. These stations were distrib- associated with mesozooplankton biomass values in uted approximately within the same latitude, in the tropical waters. To compare trends on the macroscale main water mass transport, from the northern African we used data on zooplankton sampling summarised upwelled waters to the western oligotrophic regions. from 42 expeditions by former Soviet Union research The differences between the productive (Stns 811 and vessels to the Atlantic Ocean (Fig. 5; see also Piont- 813) and oligotrophic regions (Stn 781) are shown in kovski & Ignatyev 1992). The sampling statistics are the abundance, which was 6 to 7 times greater, given in Fig. 6 and show an excellent spread through- and bioluminescence, which was about 30 times out the year. The zooplankton wet weight biomass greater (Table 2). What can also be observed is the map was created on the basis of integrated catches decrease in abundance of bioluminescent Pyrrophyta (0 to 100 m layer) conducted at 1346 stations from 1950 and copepods from the eutrophic coastal waters to 1989. To produce a map comparable with that of bio- off northwest to the oligotrophic oceanic waters. luminescence, stations with night samples only were However, it is difficult to determine from our data selected. whether or not the relative contributions of phyto- and What is obvious from the simple visual comparison of zooplankton to the bioluminescence intensity change both fields is that trends of bioluminescence, described in any systematic way in the same direction. above qualitatively, are similar to those of zooplankton For the primary production, the differences were on biomass. However, the contribution of phyto- and zoo- the order of 1 magnitude for these regions (Kon- plankton fractions to the formation of integrated biolu- dratieva & Markova 1975, Greze et al. 1984). To make minescence potential can vary over the Atlantic Ocean the spatial resolution of bioluminescence and plankton regions. To illustrate this we analysed data from a sampling comparable over depths, all values of the number of stations (Stns 811, 813, 817 and 381) variables were averaged within several layers (0-10, (Table 2, Fig. l) where coincident samples were taken 10-25, 25-50, 50-75, 75-100 m). These averaged and 3 8 Mar Ecol Prog Set 156: 33-41, 1.997

a) Monthly distribution of the Zooplankton data 1

z loo 5 50

Month

b) Year distribution of the Zooplankton data 200 180 160 g 140 .- 5 120 V) l00 LVI % 80

Zg 60 40 20 0 Fig. 6. Month and year distribution of Years the data in Fig. 5 spatially compatible data containing bioluminescence dance of bioluminescent copepods formed the main values and abundance of bioluminescent species of part of the first principal component in the highly pro- phyto- and zooplankton were subjected to principal ductive waters, whereas the bioluminescent Pyrro- component analysis and the Pearson correlation analy- phyta algae and the relatively bioluminescent sis. The first 2 principal components explained about species formed the main contribution to the second 75 % of the total variance for the given regions. The principal component which explained 37 % of the total bioluminescence potential, seston biomass and abun- variance.

Table 2. Characteristics of the planktonic community (0 to 100 m).Biolum (10-'' W cm-2].b~oluminescence intensity; Seston (mg m-'): all plankton components collected by Juday net, 36 cm mouth diameter, 124 to 145 pm mesh; Pyr,,, (103 cells m-": total abundance of Pyrrophyta algae; Pyr,,, (103 cells m-": abundance of bioluminescent species of Pyrrophyta algae. Cop. 1 (ind. m'": abundance of bioluminescent species of copepods; Cop. 2 (ind. m-3):abundance of relatively bioluminescent species of copepods; Cop. 1+2 (ind m-3):sum of the definitively bioluminescent and relatively bioluminescent species of copepods

( ~tn Blolum. Seston Pyrtot PY~IU" Cop. 1 Cop. 2 Cop. 1+2 1 Piontkovski et a1 The biolumlnesc:ent field of the Atlantic Ocean 3 9

In the low productive area (Stn 781) the first 2 princi- Ocean through the year in both hemispheres. In the pal components explained approximately equal parts northern tropical and subtropical latitudes the Canary of the total variance (31 and 34%). The relatively bio- Current passing along the northeastern African coast luminescent copepods and the bioluminescent poten- entrains the highly productive waters fed by intensive tial itself form the main contr~butionto the first princi- coastal upwelling (Herbland et al. 1983). The Canary pal component, whereas the second one is formed Current forms and feeds, In turn, the limbs of the North mainly by the seston biomass. Passat Current crossing the Atlantic approximately at latitudes 10" to 13'W (Martinenko 1988, 1990). The Interpassat Counter-Current acts in the opposite direc- DISCUSSION tion, ensuring the easterly directed water mass trans- port. The interaction of the above currents and the The map in Fig 3 represents the seasonally aver- longitudinally elongated equatorial divergence zone aged 2-dimensional field of the biolum~nescence affects the formation of enhanced values of b~olumi- potential. This type of averaging smoothes the other nescence and zooplankton b~omassand the zonally patterns and components of spatial-temporal variabil- or~entedposition of their isolines. The macroscale sys- ity. However, our goal was to create, at thls stage, the ten1 of currents in the upper layer also includes a vertl- average map of bioluminescence, to compare with cal component, which forms as the consequence of the maps of primary production and phyto- and zooplank- divergence of horizontal flows. The -driven sur- ton (Bogorov et al. 1968, Koblentz-Mlshke 1977, Sem- face divergence and the subsurface structure of water ina 1977). In this respect we wanted to complete the masses are closely coupled at the tropical Atlantic lati- mapping of general patterns of the biological structure tudes (Levitus 1982, 1988, McClain & Firestone 1993). of the Atlantic Ocean pelagia on a basin scale and to As a result of this coupling the coastal upwelling compare them later with data obtained from the events, interacting with the Ekman transport, affect deployment of 'SALPA' in the Mediterranean Basin spatial patterns of planktonic fields on an ocean basin (Bitukov et al. 1997) and with data obtained in the scale. Pacific Ocean (Gitelzon et al. 1992) In the southern hemisphere the system of currents The contr~butionof the phyto- and zooplankton frac- forms the anticyclonic gyre acting on a scale of the tions to the formation of the bioluminescence potential whole southern Atlantic Ocean (Greze et al. 1984). is controversial. The ratio and the role of components Currents propagating along the African continent, i.e. can be distinctly different over the Atlantic Ocean, the , carry the upwelled productive even within the tropical latitudes. In the northwest waters. Currents acting along the east coast of South tropical Atlantic (e.g. 3 macroscale transects of the America, i.e. the Brazilian Current, are less productive. Brazilian coast, 2" to l?" N; Fig. 1) the dominant super- The central part of the South Atlantic Ocean more or fic~albioluminescents were algae of the genera Gleno- less coincides with the position of the center of this dinium, Peridinium and Exuviaella (Evstigneev et al. anticyclonic gyre, where the most oligotrophic waters 1990). The contribution of phyto- and zooplankton are observed. For instance, differences between the fractions to the formation of the bioluminesceilce Benguela and Brazilian Currents amount to 5-10 times potential was not assessed by the authors. for the bioluminescence potential and In the upper 200 m of the the majority b~omass,5 times for phosphorus concentration and 1.5 of the stimulated bloluminescence is produced by crus- times for the averaged temperature in the upper 100 m taceans, lar\iaceans and colonial radiolai-ians, and layer (Gi-ezeet al. 1984). produced 5 to 30°,#of the measured Our observations from the tropical plankton commu- b~oluminescence (Swift et al. 1983, 1985). A cons~der- nity of the Atlantic Ocean (where the majority of our able role of zooplankton in the formation of the stimu- bioluminescence measurements were taken) are less lated bioluminescence was noted for a number of other subject to seasonal changes than at other latitudes tropical regions, e.g. the Red Sea (Rudjakov & Voron- (Greze et al. 1984). Thus, Oudot & Morin (1987) found lna 1967) and the whole tropical Pacific Ocean (Gitel- no considerable changes in the chlorophyll values zon et al. 1992). At the same time it was noted that for within the euphotic zone (5" N to 9" S, 4" W) in July regions such as the Red Sea and the Gulf of Aden both and August compared to January and February in fractions (Le, phyto- and zooplankton) contribute to the 1983. Primary production n~easuren~ents,integrated variance of bioluminescence potential during the sum- over the euphotic zone, also exhibited no remarkable mer months (Rudjakov 1968). seasonal changes in the equatorial zone (Herbland et The macroscale changes of bioluminescence inten- al. 1983). However, maps of recon- sity become understandable from a comparison with structed from data exhibit well-defined sea- the system of currents acting in the tropical Atlantic sonality (Longhurst 1993). On the basis of 1979/80 data 40 Mar Ecol Prog Ser 156: 33-4 1, 1997

it was shown that a major bloom starts in June, extends Considering the ocean basin scale, then the evalu- westwards (to 40" W) by September and regresses in ated bioluminescent field of the Atlantic Ocean is in January. sufficient concordance with the known patterns of dis- Where plankton and bioluminescence measure- tribution of zooplankton biomass. Further efforts are ments were conducted simultaneously (Table 2, Fig. 2) required to amalgamate data on zooplankton, phyto- it seems that the variance of bioluminescence itself plankton, chlorophyll a and irradiance parameters on might be high both horizontally and vertically. These an ocean scale. These data, together with temperature variations can be statistically linked with variations of profiles, would enable the bioluminescent field of the abundance of phyto- and zooplankton. For example, ocean to be derived. Algorithms relating the above- the correlation value (r) between bioluminescence mentioned parameters are being developed (Kiefer & potential and the abundance of bioluminescent species Ondercin 1993, Ondercin et al. 1996) and are the aims of copepods was 0.85 to 0.99 for Stns 81 1 and 813 from of our further studies. Such developments would the highly productive waters. The correlation coeffi- enable the bioluminescent field of the ocean to be cient between bioluminescence potential and the linked with remotely sensed physical fields on a global abundance of bioluminescent species of Pyrrophyta scale. algae was 0.78 for Stn 813. Both plankton fractions play a statistically significant role in the formation of Acknowledgements. Wethank B. Sokolov and V Vasilenko for their help in the database formation. Plankton samples the contribution to the 2 principal components which were processed by T Narusevich, L. Hlyistova and A. explain about 75% of the total variance in these Ivanova. This work was a part of the internationalcooperation regions. programme between IBSS, PML and USC and was partly In the oligotrophic waters of the western regions the funded by ONR grant No. N00014-95-1-0089. weak biolurninescence potential was not correlated with changes of Pyrrophyta algae and exhibited less LITERATURE CITED developed correlation (r = 0.6, p = 0.01) with copepods also being abundant. Artamonov YuV, Polonsky AB, Pereyaslavsky M (1987) Inves- The longitudinal/latitudinal orientation and the tigations of the macroscale water circulation in the north- eastern part of the tropical Atlantic. Deposited in VINITI elongation of spatial gradients were developed not Archives No. 391-B 87 Mar Hydrophys Inst, Sevastopol only in the bioluminescence field of the tropical (~nRussian) Atlantic but in the relationship between zooplankton Bitukov EP, Tokarev YuN, P~ontkovskiSA, Vasilenko VI. biomass and temperature (Fig. 7 in Piontkovski & Williams R, Sokolov BC (1997)The bioluminescent field as Williams 1995). The statistical relationship of the vari- an indicator of the spatial structure of the planktonic com- munity of the Basin. In: Hastings JW, ables of zooplankton and temperature in 2-dimen- Kncka U, Stanley PE (eds) Bioluminescence and chemi- sional space has the form of an ellipse, stretched along luminescence. Molecular reporting with photons. John the longitude. The ratio of the longitudinal to latitudi- Wiley & Sons, Chichester,p 169-171 nal axis of this ellipse is close to 3:l (Piontkovski & Bogorov VG, Vinogradov ME, Voronina NM, Kanaeva IP, Syetova 1A (1968)The distribution of zooplankton bio- Williams 1995). This ratio might serve as a measure of mass in the upper layer of the world's ocean. Rep USSR the spatial anisotropy of a parameter field. It was sug- Acad Sci (DokladyiAcadem~i Nauk SSSR) 182:232-245 (~n gested that the given anisotropy of the relationship Russian) between the zooplankton and temperature is the con- Chromov NS (1986)Some peculiarities of the plankton quan- sequence of the longitudinal orientation of currents titative distribution in the Atlantic Ocean. In: Biological resources of the Atlantic Ocean (Biologicheskie resyrsyl and macroscale dynamic events (convergence and Atlanticheskogo okeana),Nauka, Moscow, p 157-115 (in divergence zones) throughout the studied area. Russian] In the case of the statistical structure of the biolumi- Clarke C;L,Conover RJ, David CN, Nicol JA (1962)Compara- nescent field the stochastic component possibly points tlve studies of luminescence incopepods and other marine . Limnol Oceanogr 4: 163- 180 out the dual origin of spatially related fluctuations. Cressie NAC (1991) Statistics for spatial data John Wiley & Firstly, it is the impact of the easterly and westerly ori- Sons, New York ented currents, and secondly, it is the equatorially ori- Evstigneev PV, Bitukov EP (1990) Bioluminescenceof marine ented physical processes. They overlap and lead to the copepods (Bioluminescenciya morskihcopepod). Naukova 'cross form' of the 2-dimensional spatial autocorrela- Dumka, Kiev (in Russian) Evstigneev PV, Ritukov EP, Okolodkov JE (1993) Species tion function of the stochastic component of the biolu- composition and specificity of bioluminescence of Dino- minescent field (Fig. 4). Spatially correlated fluctua- phyceae algae. Bot Zh 78:l-15 (in Russian) tions of bioluminescence were oriented along the main Evstigneev PV, Voronova OK, Scherbatenko PV, Bocharova directions of water mass transport or macroscale equa- RK (1990) A study of biolurninescence of the superficial layer of the tropical Atlantic Ocean. Ekolo Mor 34:15-21 torial divergences and were related on a scale of 600 to (m Russian) 800 km. Gltelzon 11,Levin LA, Utyshev RN. Cherepanov OA, Chugunov Piontkovski et al.. The biolumin escent field of the Atlantic Ocean 4 1

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This article was submitted to the edltor iManuscr~ptreceived: February 28, 1997 Revised version accepted: August 8, 1997