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Radiolarian vertical distribution patterns across the Southern California current
Pergamon
Deep-Sea Research I, Vol. 42, No. 2, pp. 191-231. 1995 Copyright 0 1995 Elsevier Science Ltd Printed in Great Britain. All rights reserved 0967437195 $9.50 + 0.00
0967-0637(94)00038-7
Radiolarian vertical distribution patterns across the southern California Current STANLEY A. KLING* (Received
29 August
and DEMETRIO BOLTOVSKOY~
1993; in revisedform
5 July 1994; accepted
8 July
1994)
Abstract-Polycystine radiolarians were identified in 36 plankton samples collected at depths ranging from 0 to 2000 m at four stations extending west from about the U.S.-Mexico border (approx. 32”N, 117”W to 124”W), in November-December 1977. In total, 136radiolarian taxa were recorded, but 90% of all individuals were accounted for by only 40 of these. Highest abundances were found either at the surface, or at 25-50 m. Based on maxima in the vertical profiles of the most abundant radiolarians, three major depth-intervals were defined in the upper 300 m: O-50 m, 100 m and 200-300 m. Between-station-similarities in the specific makeups of these layers, however, were low. Thirty-nine taxa had peak abundances below 300 m at one or more stations; 11 of these are probably deep-water forms. Although in terms of individuals per liter of water filtered, upper-layer taxa are noticeably more abundant than deep species, the latter have much more extended depthranges, which might significantly enhance their sedimentary output. The inshore and oceanic stations shared very similar, warmer-water radiolarian assemblages in the uppermost 25 m, whereas the intermediate station was dominated by colder-water forms at those depths. Below 50 m, however, the inshore station had enhanced proportions of deeper- and colder-water species, differing strongly from the oceanic site. We suggest that this pattern results from circulation of the Southern California Eddy, which transports Central Water from the oceanic station on the western edge of the California Current around the intermediate stations to the inshore stations. The coldwater signal at subsurface layers of the inshore station could be reinforced by coastal upwelling and southward transport by the California Current, thus further enhancing the abundances of deeperwater radioiarians at this site. Analyses of the effect of such vertical patterns on paleoceanographic interpretations stress the importance of the signal of “environmentally neutral” deep-living species, as well as that of shells produced in the near-surface layers of distant areas and transported at depth to the region of the study.
INTRODUCTION THE overwhelming
majority of information on polycystine biogeography has been derived from surface sediment samples, whereas data on the distribution of live assemblages, and especially on species-specific vertical distribution patterns, are very scarce and largely restricted to a few oceanic areas (e.g. RENZ, 1976; MCMILLEN and CASEY, 1978; KLING, 1979; MORLEY and STEPIEN, 1984; DWORETZKY and MORLEY, 1987). Such information, however, has an important bearing on current problems associated with the fate of organic matter generated in the upper mixed layer (e.g. GOWING and GARRISON, 1992). Analyses of vertical distribution patterns can also yield valuable data on the modes of sedimentation
“416 Shore View Lane, Leucadia, CA 92024, U.S.A. ?Departamento de Ciencias Biologicas, Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires, 1428 Buenos Aires, and CONICET, Argentina. 191
192
S. A. KLINC
and D. BOLTOVSKOY
of radiolarian shells and help assess the relative importance of dissolution en route to the sea-floor and of destruction by grazing (MILLIMAN and TAKAHASHI, in press; BOLTOVSKOY and ALDER, 1992a). Many paleoceanographic studies based on comparisons between surface sediment distributions and downcore sequences employ analytical techniques (e.g. factor analysis and transfer function techniques: IMBRIE and KIPP, 1971) that assume irrelevance of living organisms in the overlying plankton. However, these assumptions do not take into account biases associated with vertical distribution patterns of such deepliving groups as radiolarians. In addition, at any given locale bottom thanatocoenoses are made up of locally produced shells and, to different extents, by shells produced elsewhere and transported laterally via subsurface and deep currents (BURCKLE, 1981; POKRAS and MOLFINO, 1986; BOLTOVSKOY, 1988, 1991, 1992; BOLTOVSKOY and ALDER, 1992a). Thus, climatic oscillations which affect the strength of these currents (and, consequently, the intensity of advection of sedimenting shells) can introduce bias into direct comparisons between recent and past geographic distribution patterns. The present work describes the vertical (O-2000 m) distribution of polycystine Radiolaria at three closely spaced sites off the Southern California coast dominated by a strong gradient in physical and biological variables. In addition, duplicate sampling at one of the sites provides information on the temporal (approx. one week) variability in polycystine makeup and distribution. Although the patterns analysed are but a snapshot image of a highly variable situation, they nevertheless offer clues for the interpretation of the ecological settings of the radiolarians involved and, by extension, on their paleoceanographic significance in the fossil record. MATERIALS
AND
METHODS
Samples were collected from a line of stations extending west from about the U.S.Mexico border (Fig. l), as part of BMET Cruise in late November and early December of 1977. Station data are as follows: Sta. A: 23 Nov. 1977,32”38’N, 117”29’W; Sta. Cl: 25-27 Nov. 1977,32”30’N, 120”26’W; Sta. C2: 30 Nov.-3 Dec. 1977,32”38.25’N, 120”35.75’W; and Sta. F: 5-6 Dec. 1977,32”38.23’N, 123”53.96’W. Field and laboratory methods for this study are essentially the same as those described by KLING (1979). Plankton samples were obtained with opening-and-closing nets controlled and monitored by an acoustically operated system described by BAKER et al. (1973). The nets, made from Nitex nylon mesh with 62-pm opening, were designed to present a 1 m by 1 m cross section to the towing path while being towed horizontally with the opening frame at a nominal 45 degree angle. They were about 3.5 m long midway along the tilted vertical frame members. Flow-meters were not available during the tows reported here, so volumes of water filtered were estimated on the basis of distance towed, calculated from ship speed and time, and net mouth diameter (1 m2). Earlier experience with the same net system equipped with an external flow meter indicates that distance calculated from ship speed and time is a reliable measure of distance travelled. Correlation between metered and calculated distances were very highly significant (Y = 0.999, P < O.OOl), which suggests that our distance estimates are generally correct. Nevertheless, this ignores the effects of net clogging, which are admittedly serious despite the relatively large size of the nets (1 m2 x 3.5 m) and the mesh openings (62,um). To compensate in some way for this shortcoming, the length of tows was much shorter in
Radiolarian vertical distribution patterns
193
the upper, plankton-rich layers (on the order of 100 m at 0 m, 200 m at 25 m) than those in deep waters, where particulate materials are scarce (700 m at 500 m, 1500 m at 1000 m, 3000 m at 2000 m). This does not ensure that the values given are correct, but it does lend support to the contention that they are at least internally consistent. We have attempted to minimize the effects of water volume inaccuracies upon the reported analyses in two ways. First, where we used numbers of individuals per cubic meter, for example to identify abundance maxima (Table 2), we considered only values that exceed background levels by about five times or more (see the vertical profiles summarized in Fig. 7). Second, most of the statistical calculations were based on data transformed into percentages. In the laboratory, samples (preserved in formaldehyde) were split and one fraction combusted to rid it of entangling soft tissue. Measured subsamples of the combustion residues were transformed to cover glasses and dried. Cover glasses were mounted on glass slides with Pleurax. All specimens on a cover glass were identified and counted. Unidentified forms were lumped in a separate category. Counts in all categories were converted to numbers per m3 of water filtered. Unidentified radiolarians, including juveniles and incomplete shells, made up very large proportions of the overall assemblages retrieved: 30.7-66.7% (see Table 1). These individuals were only included in analyses of total radiolarian standing stocks. Speciesspecific distributional interpretations, especially those based on percentage data, are obviously affected by these high proportions of unidentifieds, yet the bias seems to be evenly distributed over the entire database, since no relationship was found between the percentages of unidentifieds and site or depth. The number of individuals per sample scanned varied between 156 and 2409 (Table 1). Despite this wide range in sample-size, the database seems to represent adequately the overall specific diversity of the area insofar as no correlation was found between the numbers of specimens identified and the numbers of species recorded (1. = 0.173, P > 0.1). OCEANOGRAPHIC
SETTING
The transect of stations for this study extends across the California Current (REID et al., 1958) from the Southern California Borderland to the deep Pacific basin (Fig. 1). Station A was over the San Diego Trough with a bottom depth of about 1000 m. Stations Cl and C2 were located just west of the Patton Escarpment where water depths exceed 4000 m. Station F was in the deep ocean where water depths are in the 5000-m range. Hydrographic casts were made at each station and the resulting O-300 m temperature and salinity profiles are presented in Fig. 2. Because of the equipment failures, no salinity data are available for Stas C2 and F. The temperature profiles indicate that the surface mixed layer extends to a depth of about 100 m at the western Sta. F, shallows to about 50 m at the central Sta. C2 and 25 m at Sta. Cl, and shallows further to or near the surface at the inshore Sta. A. Salinity profiles are marked by a shallow salinity minimum at about 50 m at Sta. Cl and A (where data are available). A shallow salinity minimum is a regional feature of the California Current (REID et al., 1958;REID, 1973). The deeper salinity minimum of the North Pacific Intermediate Water (REID, 1965) is typically not maintained in the California Current, and does not appear in these profiles. The major component of flow in the California Current is southward, However, a subsurface countercurrent is also a prominent feature, and at times the countercurrent
194
S. A.
KLING
and D.
BULTOVSKOY
Table 1. General data for the materials surveyed. Equitability (E = HE&,,,, = @(Pi.logz Pi) + Ps.logz Ps); where Pi is the proportion of the ith species, and s is the total number of species in the sample; PIELOU, 1966), is inversely proportional to the degree of numerical dominance of one or a few species over the rest of the assemblage, whereas specific diversity (H’ = Z Pi .log2 Pi, where Pi is the proportion of the ith species; SHANNON and WEAVER, 1949)) increases with increasing equitability and species richness
station
Total individuals
Depth
bd.
m-3
counted
I ,4
300-1000
Specific diversity
Equitability
No. of spe
1661 966 521 608 111 121 64 37 9 21
42.45 40.27 40.56 52.98 56.97 49.86 54.14 53.23 51.47 46.75
4.734 4.544 4.544 4.771 4.289 5.023 4.881 4.983 4.460 5.146
0.786 0.801 0.790 0.792 0.790 0.854 0.837 0.834 0.863 0.842
65 51 54 65 43 59 57 63 36 69
665
47.18
4.651
0.8M
56
588
50
51.09
4.899
0.846
57
50 UN 200
156 700 910 1227 596
108 240 416 401 102
66.67 54.00 35.82 60.15 44.20
4.224 4.171 4.383 3.843 4.632
0.899 0.801 0.789 0.692 0.791
26 37 47 47 58
300 400 500 750 1000 2000
706 284 328 286 291 316
121 31 38 20 11 8
42.41 44.01 41.46 40.21 51.55 49.05
5.042 5.155 5.007 5.013 5.068 4.777
0.831 0.909 0.870 0.871 0.893 0.875
67 51 54 54 51 - 44
716
231
so.54
4.382
0.800
47
379
44
43.93
5.057
0.875
55
304
9
50.30
4.923
0.884
48
622 1843 1925 1673 2347
429 633 440 96 403
49.04 50.03 52.47 64.67 48.53
4.319 4.494 4.772 4.602 5.241
0.812 0.746 0.795 0.782 0.816
40 65 64 59 86
594 782 1311 319 1308 275
102 89 150 26 112 8
42.09 63.94 52.56 30.72 50.42 44.00
4.732 4.353 5.037 3.466 5.209 4.834
0.804 0.771 0.814 0.643 0.842 0.880
59 50 73 42 73 45
1501
351
51.14
4.693
0.793
62
863
96
47.94
4.559
0.775
59
792
60
47.21
5.022
0.861
59
822
1134
44.18
4.559
0.812
49
1003 1751 766
459 573 125
51.77 42.21 37.91
5.031 4.745 5.067
0.845 0.762 0.816
62 75 74
1086
573
44.02
4.850
0.809
65
m mean
O-300 m mean 300-1000 m mean 1000-2000 m mean
300-1000
bldh.
2409 1407 1140 1772 646 706 556 650 307 723
0 2s
c2
% unid.
m mean
Radiolarian
vertical
125
distribution
195
patterns
120'
115 35"
32"
Fig.
1.
Fig. 2.
Sampling
Vertical
locations
and November (O-500 db),
Temperature
(“C)
10
15
O-300
195C1964 after WYLL~E
mean geostrophic (1966).
flow
Salinity
to\-)
m profiles of temperature and salinity at the stations shows profiles down to the deepest layers covered.
surveyed.
at the surface
Inset
graph
reaches the surface. The surface countercurrent is particularly well developed close the coast in southern California and in winter months. These opposing currents define a semipermanent circulation system (the Southern California Eddy) over the Borderland in the broad embayment of the southern California coastline south of Point Conception to Punta Baja in Baja California (the Southern California Bight). As a result, the central part of the region may differ hydrographically (e.g. lower temperatures and salinities) from the waters either farther onshore or offshore. Similarly, inshore waters may be more closely
196
S. A. KLING
and D. BOLTOVSKOY
related to the far offshore Central Water Mass than to the core of the California Current. We note in our data that surface temperatures are slightly lower at the central stations (Cl, C2) than at the outer stations (A, F) (Fig. 2). Available surface salinity data are consistent with the temperature observations: lower at Sta. Cl than at A. The California Current is a very complex system that changes rapidly in space and time. However, PELAEZ and MCGOWAN (19S6) recognized a number of major features in the southern California Current that.appear to have both hydrographic and biologic significance. The structures were identified from satellite images of phytoplankton pigment concentrations near the ocean surface. Three features are of interest to this study. An area of high pigment concentration is probably related to the Southern California Eddy and located over the bathymetric high area of the outer Borderland basin system (the location of Stas Cl and C2). Secondly, a low-pigment intrusion flows from the south in the countercurrent close to the coast off San Diego in the region over the deep San Diego Trough (Sta. A; see Fig. 2), and is especially well-developed in summer and fall months (July-December). This low-pigment water is related to a third feature, the low-pigment, oligotrophic waters of theNorth Pacific Cental Gyre to the west (Sta. F), which circulate around the southern and eastern borders of the high-pigment area. Although in detail these patterns shift position, expand and contract, and change in pigment concentrations with time, they were always identifiable in some form. A seasonal cycle in their development noted by PELAEZ and MCGOWAN (1986) were repeated in the three years’ records they analysed. RESULTS AND DISCUSSION Specific makeup
In total, 136 radiolarian taxa were identified in the collection (see Appendices 1,2). This number compares favourably with previous studies of complete polycystine assemblages in the California Borderland. BOLTOVSKOY and RIEDEL (1987) recorded 155 species in forty-eight O-100 m plankton tows along the California Current; and CASEY (1966) reported approx. 200 species from the same area between October 1962 and February 1964. However, as in previous studies, a rather small proportion of these 136 taxa was represented by common to abundant forms, with the bulk being recorded sporadically and in very low numbers. Thus, when the entire database is considered, only about 40 radiolarians accounted for ca 90% of all identified individuals. Although the rare species do convey important ecological information, probably even more than the abundant ones (e.g. BOLTOVSKOY, 1989), their scarcity poses serious methodological problems to interpretation of associated environmental signals. In fact, absences of these rare organisms from the samples are questionable, rather than presences, because of the impossibility of distinguishing the combined effects of ecological exclusion and insufficient sample-size. The only practical way to overcome this problem is to restrict the database to those species whose absences can be reasonably assumed to result from environment-related causes (rather than to procedural artifacts). Therefore, we based many of our subsequent detailed analyses on a reduced set of 41 taxa with at least 20 individuals in one or more samples. This arbitrary cutoff value seems an acceptable compromise which retains a moderately large inventory, yet excludes the questionable absences. These 41 abundant radiolarians comprise 6491% of the entire inventory of each sample (excluding the
Radiolarian
vertical
distribution
197
patterns
‘50 100 300
n
cl
Fig.
3.
Vertical
distribution
of total radiolarian
concentrations.
unidentified specimens). However, in agreement with the pattern of total abundances (Fig. 3), their relative contributions were higher in the top layers than at depth.
Interpretation
of the vertical profiles: in situ living populations vs dead sedimenting shells
A major problem for the assessment of radiolarian depth ranges is the lack of information on the physiological status of the individuals (i.e. live cells or dead, sinking skeletons). It is well known that, although specimens retrieved in the uppermost layers represent mostly live organisms, the proportions of empty shells increase significantly with increasing depth (e.g. PETRUSHEVSKAYA, 1971; TAKAHASHI, 1983/1984; GOWING and COALE, 1989; BOLTOVSKOY et al., 1993a). In our materials concentrations below the uppermost maximum decrease either more or less gradually, or abruptly, but most profiles suggest a restricted preferred depth range, with a “tail” made up of “stray” live (?) specimens and settling dead individuals (Fig. 3). In some instances, however, the profiles show several maximum throughout the water-column (e.g. Sta. Cl, see Fig. 3). These patterns are amenable to alternative interpretations. Although the upper peak is probably due to live individuals, the lower ones can be: (1) enhancements due to favourable local conditions for some of the taxa concerned; (2) accumulations of skeletons transported laterally from elsewhere; (3) sinking accumulations resulting from short-lived blooms at shallower depths; (4) concentrations of sinking shells derived from discontinuities in the density profile of the water-column, or a combination of the above. The secondary peaks below primary ones higher in the water-column largely represent settling shells, rather than in situ blooms, is suggested by a close analysis of the four maxima in the profile of Sta. Cl (at 25, 200, 500 and 1000 m, cf. Fig. 3). Although the uppermost peak (at 25 m) is totally dominated (91%) by taxa that are at maximum abundance precisely in this particular stratum, the dominance of locally peaking species
198
S. A. KLINC andD. BOLTOVSKOY Ten
25 m species
Two
200
m species
Three
500
m species
Three
1000
m species 3
T. octacantha E. acuminarum P. t&bum D. tetrathalamus 5. irregularis L. quadran u/a Perrdium (?P sp. P. praetextum T. trachelium P. hertwigii
tetras
1800 1 zooo-
davisiana
L. buetschli cornutoide
II
0
80
160
10
20
30
Individuals
L
Spongorus sp. T. bicornis f?) sp. 1
Carpocanium
Dicryophimus
per
0
1
sp. 2 2
m-3
Fig. 4. Vertical profiles of the species characteristic of the four peaks in total radiolarian abundance at Sta. Cl (cf. Fig. 3). In proportion of total individuals, these taxa were 22 times more abundant at the given layer than at any of the other three layers compared.
decreases to 63% at 200 m, 20% at 500 m and 25% at 1000 m. Conversely, the contribution of species presumably generated elsewhere (i.e. peaking above a given depth), is 32% at 200 m, 66% at 500 m and 75% at 1000 m. The remainders, however, are made up of taxa that are at maximum abundance within the given layer, and therefore probably represent live populations, or at least assemblages not generated in the upper strata of the same locale (see below). Figure 4, based on a selection of the most reliable data points, shows that the uppermost (25 m) maximum at Sta. Cl is dominated by 10 clearly surface-dwelling radiolarians (52% of the overall assemblage at this depth). These taxa account for 14,6 and 10% at 200,500 and 1000 m, respectively. Theocalyptra davisiana cornutoides and Larcopyle buetschlii account for 19% of all identified radiolarians at 200 m, although they represent only 1% at 25 m, and 4-5% at 5OGlOOO m. Spongurus sp., Theocalyptra bicornis and Dictyophimus sp. 1 comprise 17% of the (identified) individuals at 500 m, whereas they are absent altogether at 25 m, and represent 0.7 and 4.5% of the radiolarians at 200 and 1000 m, respectively. Finally, Siphocampe arachnea, Porodiscus rnicroporus and Carpocanium sp. 2 are noticeably more abundant at 1000 m than elsewhere. Although their overall contribution to this layer is a low 4%) they represent only O-0.6% of the assemblages at 25, 200 and 500 m. Figure 4 also shows that minor yet noticeable peaks for the taxa characteristic of the strata reviewed also occur at some of the levels of maximum total radiolarian abundance (cf. Fig. 3). Th us, the ten 25 m species also have high concentrations at 200 m; the two 200 m ones at 25 and 500 m, etc. Although secondary maxima above the layer of maximum abundance are most probably due to live individuals (probably the initial part of the downward abundance increase), it seems difficult to envision that a species that thrives, for example, in the upper 50 m can again find very favorable conditions at 200-1000 m. The percentages of Fig. 4 are based on total radiolarian absolute abundances, for which our
Radiolarian
vertical
distribution
199
patterns
Total species recorded
Species added (Oh of total)
Fig.
5.
Total number new species
of radiolarian species added to the inventory
recorded in each sample (top panels), and numbers of the overlying samples (bottom panels).
of
estimates are somewhat rough (see Materials and Methods). Thus, biases in these numbers, on which concentrations of individual species are based, greatly affect the shapes of the profiles shown. That radiolarian assemblages at depth increasingly represent flux of skeletons out of the upper layers is also strongly supported by the fact that neither the number of taxa, nor specific diversity or equitability, vary systematically with depth (Table 1). Figure 5 illustrates the numbers of species recorded in each sample (top panel), and the new species added to the total inventory of each station as one progresses from top to bottom (bottom panel). It is noteworthy that, although most of the taxa are present at all depths, deeper samples contribute successively smaller numbers of new taxa to the roster encountered in the overlying waters. The upper 50 m account for 55-72% of the total diversity of each site (Fig. 5). Interestingly, the higher vale (72%) is that of Sta. Cl, which showed the most pronounced subsurface abundance peaks (Fig. 3). This reinforces the above-discussed premise that these deep maxima at 200,500 and 1000 m are either biases in our estimates of total radiolarian abundance, or accumulations of dead skeletons.
200
S.A.KLING~II~D.BOLTOVSKOY %
of
“live”
Polycystines
“Live” polycystines (regression vs. log depth, r = - 0.897) % of “live” polycystines (actual data points)
-llIlTota’ 5,
polycystines per m-’
polycystines
per
0
Total
Fig. 6. Average percentage of “live” polycystines and mean total radiolarian shells vs sample depth;
m-3
(see text for description of assumptions used) based on combined data for entire collection.
We also attempted to address the problem of the differentiation between living and dead radiolarians by assuming that individuals retrieved at and above the level of a species’ maximum abundance were live cells, and that all those below the maximum were dead skeletons settling through the water column. For all taxa “live” and “dead” specimens were summed separately, and the average ratio between the two figures was calculated using all four sites samples. Figure 6 illustrates the curve thus obtained, superimposed on the regression of percentage of “live” individuals on log depth (meters + 1). The assumptions used in this approach have some obvious pitfalls: (1) a species can peak at more than one depth-horizon; (2) downward extensions of the preferred living depth-range are ignored and attributed to settling shells; and (3) local accumulations of sinking skeletons can be mistakenly taken as preferred living depths. Nevertheless, interestingly, these results agree remarkably well with a similar analysis of the vertical distribution of live and dead radiolarians based on the presence of protoplasm in their shells. In the equatorial Atlantic about 2% of the polycystines collected in sediment traps at 850 m are individuals with protoplasm (BOLTOVSKOY et aE., 1993a). It should be emphasized that differentiation between skeletons with and without protoplasm in vertical plankton profiles can furnish clues on the living depth ranges of the species, yet it does not provide unequivocal information because of the uncertainties associated with the speed of decomposition of the protists’ cytoplasm. BOLTOVSKOY and LENA (1970)) for example, concluded that specimens of several planktonic Foraminifera
Radiolarian vertical distribution patterns
201
still contained protoplasm in their shell 98 days after death. This lapse is significantly longer than the time it takes a radiolarian shell to reach the sea-floor (TAKAHASHI and HONJO, 1983). Information on living depth ranges vs export ranges are also intimately linked with processes responsible for the production of new shells, and with the sinking speeds of the skeletons, For surface-dwelling taxa, for example, comparison of standing stocks in the upper layers with concentrations at depth can furnish hints on reproduction rates. In our database, the mean abundance of the 20 radiolarians which peaked at 0 m at Sta. F (see below) was approx. 346 ind m -3 in the O-100 m stratum (which, presumably, is where the majority of these species reside). Between approx. 600 and 1000 m their concentration was rather stable, averaging 3.4 ind. rnm3. Thus, assuming a sinking speed of 1.50 m day-’ (TAKAHASHI and HONJO, 1983), losses by sinking from O-100 m, as recorded by our 60s 1000 m samples, are 1.47% of the population at O-100 m. This translates into a turnover rate of 68 days. This estimate is somewhat higher than previous (indirect) estimates of radiolarian reproduction rates, which suggest values ranging around 20-30 days (see review in CARON and SWANBERG, 1990). Faster sinking speeds and shallower living ranges for the taxa considered, among other factors, would increase this doubling rate. A major point, however, is most probably the loss of shells due to destructive grazing by larger zooplankters. BOLTOVSKOY and ALDER (1992a), for example, concluded that over 90% of the radiolarian shells produced at O-400 m in the Weddell Sea are destroyed beyond recognition above 900 m. Our results suggest that, if radiolarian life-spans are effectively around 20-30 days, in the area of the study destructive grazing eliminates 30-50% of the output from the upper 100 m. It should be stressed that the impact of this process is likely quite uneven on radiolarian taxocoenoses, more fragile skeletons being eliminated in larger percentages than the more robust ones. Comparison of sediment trap vs surface sediment assemblages in the eastern equatorial Atlantic (BOLTOVSKOY et al., 1993b) suggest that the relative contribution of spongodiscid shells might be over-represented in the “grazed assemblages”, whereas the nassellarians, and especially their young, developing forms, might be under-represented. Based on the above considerations, our analyses of the vertical distribution patterns and depth preferences of individual radiolarian species paid special attention to the uppermost layers of peak abundance of each taxon. Vertical distribution patterns Highest radiolarian abundances were recorded either at the surface (Stas A, F), or at 25 (Sta. Cl) to 50 m (Sta. C2). Secondary maxima below this upper layer were generally minor, with the exception of Sta. Cl, where samples from 200 m, and, to a lesser extent, from 500 to 1000 m, presented noticeable peaks in radiolarian concentration (Fig. 3 and Table 1). Because most of the radiolarians were concentrated in the upper most layers, and in order to facilitate comparisons between Sta. A (which was sampled to 300 m only) and the rest of the sites surveyed, detailed treatment of vertical patterns was subdivided into two depth-intervals: O-300 m, and O-2000 m (for the species which showed major peaks below 300 m only). In the O-300 m section a total of 38 species was included (of the 41 for which over 20 specimens were recorded in at least one sample, Spongopyle osculosa, Larcopyle
202
S. A.
KLING
and D.
BOLTOVSKOY
buetschlii and Spongurus sp. were eliminated because they had deep distribution patterns). In the O-2000 m description we considered all the taxa that peaked below 300 m in one or more sites. The upper 300 m
In general terms, species-specific vertical profiles varied widely between stations. As a first approach to the assessment of congruence between sites, we correlated all station pairs of vertical O-300 m profiles of the 41 most abundant radiolarians. Only 16% of the indices thus obtained were significant (positive) at the 95% level, and about twice as many (33%) were negative. Surprisingly, the lowest proportion of significant (P < 0.05) correlations was yielded by the two series carried out at the same locale (Cl-C2: 7%), while highest similarities were those between Stas A and F (32% of the indices significant at P < 0.05). A more in-depth analysis based on curve-fitting by eye and cluster analysis (not shown), suggested several species groups with more or less clear preferences for a restricted depth interval. Figure 7 and Table 2 summarize these results, which are described as follows: Group I: species with abundance maxima near the surface: Subgroup 1: 0 m maximum and sharp decrease below. Subgroup 2: 0 m maximum and gradual decrease below. Subgroup 3: 25 m maximum. Subgroup 4: 50 m maximum and secondary 0 m maximum (Sta. Cl only). Subgroup 5: 50 m maximum. Subgroup 6: 0 and 100 m marked maxima. Group II: species with subsurface maxima: Subgroup 7: 100 m maximum. Group III: the deepest t&300 m species: Subgroup 8: 200 m maximum. Subgroup 9: 200 and 300 m maxima (one radiolarian only). Subgroup 10: 300 m maximum. Of these 38 radiolarians in the O-300 m data set, between 37 (Stas A and F) and 29 (Sta. Cl) were included in one or more groups, and none was excluded from all groups. Several profiles were not assigned to groups because of irregular or ambiguous vertical patterns. Similarities between the specific composition of groups I-III at the four stations were variable and generally rather low. Of the 34 species which were most abundant in the upper layers at one or more stations, only 14 entered group I at all four sites surveyed (asterisked in Table 2). Another seven entered group I at three of the four stations, and the remaining 12 only at two or one. Fifteen radiolarians had peak abundances at 100 m at one of the stations (group II), three at three stations, and only two taxa (Eucecryphalus sp. and Litharachnium tentorium) showed similar 100 m peaks at the four stations (Table 2). Finally, out of the 10 taxa which peaked at 200 or 300 m at one or more stations (group III), only Theocalyptra bicbrnis and Theocalyptra divisiana cornutoides showed coherent deep vertical patterns throughout the area (Table 2). Figure 8 summarizes similarities in radiolarians species compositions between stations for the three depth-intervals described on the basis of the vertical patterns defined. The uppermost layer is faunistically the most homogeneous throughout the area: on the average, ca 63% of the species which peak within this depth-interval are common to two or
Radiolarian
vertical
distribution
patterns
203
Fig. 7. Vertical distribution of the most abundant radiolarians in the O-300 m layer based on 2.5 m interpolated profiles. Shaded areas encompass all profiles grouped within each of the general patterns defined (see Table 2 for species included in each group).
more stations (asterisked in Table 2). Mid-depth and lower-depth taxa (groups II and III), however, vary considerably more between sites (26 and 30% in common, respectively), yet these similarities are still considerably higher than those between groups (9 to 4%) see Fig. 8). In addition to the abundant radiolarians, vertical ranges were also assessed (on the basis of maximum abundances throughout the water-column) for 28 taxa that were less abundant yet present at 23 of the four stations, and followed rather similar trends throughout the area. Table 2 illustrates the data in question: species 87 through seven (‘Upper 50 m” in Table 2) consistently peak between 0 and 50 m, and therefore have ranges similar to those described for group I. For taxa 43-127 (“O-100 m” in Table 2) and 139-52 (“25/50-100 m” in Table 2), peak abundances change from site to site between 0 and 100 m, with deepest maxima often occurring at Stas A and F. Radiolarians 129 through 28 (“100-300 m” in Table 2) center on 100-300 m. Finally, the last three taxa illustrated
204
S. A. KLING
and D. BOLTOVSKOY
Radiolarian ; . i i E i
E e : . E i : L
L .
L i
.
i
.
vertical
distribution
patterns
205
206
S. A. KLING
Percentage
and D. BOLTOVSKOY
of species
in common
3
Fig. 8. Similarities between radiolarian specific makeups for the &300 m vertical distribution groups I-III (cf. Fig. 7 and Table 2). Numbers denote species in common as a percentage of the total number of taxa encompassed by the two groups compared.
(ID nos. 9, 24 and 46, “200-300 + m” in Table 2) are consistently deep dwellers, with patterns similar to those of group III (Fig. 7). In agreement with the profiles for total shells (Fig. 3), the same radiolarian species tended to dwell shallowest in the water-column at Sta. F, and deepest at Stas Cl and C2. In the upper 300 m there did nt seem to be any relationship between preferred depth and differences in depth distribution from one station to another. In other words, most radiolarians occupied generally deeper strata at Cl-C2 than at A, and deeper at A than at F, regardless of their individual profiles. Thus, for the 13 consistently upper layer taxa (asterisked in Table 2), for example, the mean weighted population depth [WPD = C(ni . di)l%zi, where ni is the number of individuals at depth i] varied from 52 m at Sta. F, to 69 m at Sta. A, to S&91 m at Stas Cl-C2. For the rest of the 41 abundant radiolarians these values were 113, 137, 145 and 159 m, respectively (calculations based on O-300 m data only). The O-2000 m layer Thirty-nine of the 136 radiolarians identified had maximum abundance below 300 m at one or more stations (Table 3). Only three of these, however, were abundant (i.e. 220 individuals recorded in any one sample): Larcopyle buetschlii, Spongopyle osculosa and Spongurus sp. The latter two had fairly consistent patterns peaking generally at or below 300 m (Table 3, Fig. 9), but L. buetschlii’s abundances varied more between stations (Fig. 9).
Radiolarian
vertical
distribution
patterns
207
Table 3. Species with peak concentrations at depths of 300 m or more at one or more stations (selected from entire database). Data for each site are maximum abundance recorded (ind. mP3) and, in parentheses, corresponding depth. Frames highlight maxima at 300 m (thin line), and below 300 m (thick line). Last column gives maximum abundance in entire collection (ind. mP3). Taxa are sorted in descending order of maximum concentration recorded Results of ANOVA CZ/F 1 Cl/F
Total signif=ant
contrasts
1
16 1
1 Cl/C21
14 1
2 1
A/F
1 A/C2
18 1
1 A/Cl
10 1
12
S. A. KLINC and D. BOLTOVSKOY
Cyrtolagena
@
Larcopyle
buetschlii
Fig. 9 Vertical distribution
laguncula
Q
Spongopyle
osculosa
of deep-dwelling species. Numbers above profiles indicate scale maxima in ind. m-3.
Among the rare species, only Cyrtolagena laguncula and Sethoconus (?) sp. had maximum abundances below 300 m at the three sites with data from the deeper layers (Table 3, Fig. 9). In addition to the above, a few other radiolarians tended to peak below 300 m, yet their concentrations were too low to define reliable vertical trends: Botryostrobus aquilonaris, Siphocampe arachnea, Phormostichoartus corbula, Bathropyramis woodringi, Artostrobus annulatus and Sethoconus (?) sp. aff. S. (?) dogieli (cf. Table 3). Depth-abundance
relationships
The above-described depth preferences are probably to some extent related to overall abundances of the species in the entire database. Figure 10, which summarizes the stationto-station ranges in peak abundance for the 136 taxa, shows that deep-living forms are mostly located on the right-hand side of the graph (i.e. lowest mean abundances), whereas the upper-layer taxa are the most abundant. In addition, the station-to-station vertical ranges tend to be larger for the deeper, less abundant radiolarians. We suggest that this figure illustrates the combined consequences of several effects: (1) The uppermost layers host the numerically dominant species, whereas specific abundances decrease sharply at greater depths; (2) Vertical ranges tend to be better circumscribed in the upper layers, where environmental gradients with depth are steeper, than in the more homogenous meso- and bathypelagic layers. This trend has long been recognized, both for radiolarians (e.g. KLING, 1979), and for many other planktonic groups (VINOGRADOV, 1968); (3) Because deep-water species are less abundant, sample-size-related errors in the definition of their vertical ranges are larger. Thus, whereas (1) and (2) are actual relationships, (3) is an artifact.
Radiolarian vertical distribution patterns
Radiolarian
209
species
Fig. 10. Station-to-station ranges of depth of maximum concentration for all the radiolarians identified (hanging bars), as a function of their mean abundance in the entire collection (gray shading).
Comparison
with previous data
Although detailed comparisons of our results with previous studies is seriously restricted by methodological and nomenclatural differences, data summarized in Table 2 generally agree with, and amplify, previously published results. KLING'S (1979) survey of the vertical distribution patterns of polycystines of the central north Pacific is the study most closely comparable to the present one. Comparison of abundance maxima in the vertical profiles of 39 (out of total of 43 identified by KLING, 1979) radiolarians indicates that, with the exception of six taxa, vertical patterns derived from the present database encompass the ranges found by KLING (1979) in the central north Pacific. Only Cornutella profunda, Lamprocyrtis nigriniae, Theocalyptra bicornis, Botryostrobus aurituslaustralis, Dictyophimus infabricatus and Lophocorys polyacantha had deeper maxima in the central north Pacific than in any of the BMET stations. None of the above, however, exceeded 3.5% of all radiolarians in KLING'S (1979) collection or 3.7% in our database. Species-specific relative abundances, on the other hand, show differences consistent with warmer-water conditions at the INDOPAC sites (surface temperature ca 22-23”(Z), than at the sites covered in this work (see Fig. 2). Thus, in the O-100 m layer the typically warm-, oligotrophic-water radiolarians Heliodiscus asteriscus, Lithopera bacca, Phormospyris stabilis s.1. and Siphonosphaera polysiphonia were 5-165 times more abundant in KLING'S (1979) collection than off San Diego. Conversely, the BMET area O-100 m layer had higher proportions (5-50 times) of Botryostrobus aurituslaustralis, Pterocorys clausus, Spongocore cylindrica, Theocalyptra davisiana cornutoides and a few other characteristically colder water species. The BMET database is also generally consistent with the collection reported by BOLTOVSKOYand RIEDEL (1987) from O-100 m samples retrieved throughout 1972 in the California Current. Differences other than those attributable to variable sample-size and species-definition disagreements chiefly affect some of the deep-dwelling radiolarians which were not recorded in Boltovskoy and Riedel’s shallow tows (e.g. Artostrobus annulatus, Bathropyramis woodringi, Botryostrobus aquilonaris, Lamprocyrtis nigriniae, Cyrtolagena laguncula, Siphocampe arachnea, Spongopyle osculosa, and a few others; cf.
210
S. A. KLINC and D. BOLTOVSKOY
Table 2). Large dissimilarities in percentage contribution between the two collections were noticed for the deep-dwelling Theocalyptra davisiana s.1. and Spongocore cylindrica (ca lo-40 times more abundant at BMET), whereas the warmer-water Heliodiscus asteriscus, Polysolenia murrayana and Solenosphaera polyrnorpha were 8-13 times more abundant in BOLTOVSKOY and RIEDEL’S (1987) collection. Larcopyle buetschlii, however, whose maxima were at or below 100 m at all the sites surveyed (Appendices 1, 2), was approx. 20 times more abundant in the O-100 m California Current plankton than in the BMET samples. Hydrographic
control of radiolarian
patterns
Sparsity of contemporaneous hydrographic data for this study limits direct comparisons between species distributions and hydrographic patterns. Further complications arise from the complexity and instability of the California Current system, which lend importance to discrepancies in the time and spatial scale between hydrographic and plankton data collections. Nevertheless, some general patterns seem to emerge. As described above, analysis of geographic changes in radiolarian vertical distribution and assemblage makeup show that the two stations located at the extremes of the transect (A and F) are rather similar insofar as highest overall concentrations occur closest to the surface at these stations (Fig. 3), and assemblage compositions in the upper 25 m are more similar between these two locales, than between either one of them and Cl or C2 (Fig. 8). On the other hand, comparison of O-300 m relative abundances indicates that these two stations (F and A) differ most in their specific composition: of the 41 most abundant radiolarians, 21 showed significantly higher relative abundances at either Sta. F or Sta. A (Table 4 and Fig. 11). The nine species with highest values at Sta. F had, on the average, 5.1 ind. rnp3 at this site, and only slightly over 1 ind. rnp3 at the other three stations (Fig. 11). Six of these nine taxa are conspicuous members of group 1 (upper 50 m: cf. Tables 2 and 4). In contrast, species characteristic of Sta. A (12 in total, cf. Table 4) include several 100-300 m (Theocalyptra bicornis, Eucecryphalus sp, , Litharachnium tentoriurn, Dictyophimus infabricatus, D. gracilipes), and deep-water radiolarians (Spongurus sp.) (cf. Tables 2,3 and 4). In terms of the overall assemblages, differences between Stas A and F are quite significant insofar as they affect approximately half of the radiolarian shells recorded at Sta. F, and about 35% at Sta. A. This seemingly contradictory result is consistent with the regional circulation pattern as described above. We note that Central Water impinging on the western edge of the California Current, where Sta. F is located, is carried by the Southern California Eddy around Stas Cl and C2 and then north along the coast to the area of Sta. A (see “Oceanographic setting”). Since this Central Water is warmer than typical California Current waters, it overlies the latter near the coast. This may account for higher 0 m temperatures at Stas F and A. However, a thermally uniform 75 m upper mixed layer characterizes F, whereas temperatures drop sharply from the very top of the water-column at A (Fig. 2). This hydrographic pattern is consistent with radiolarian distributions. In the O-25 m layer of the area surveyed the 12 species which peak at the surface and drop sharply thereafter at Sta. A (group A-l in Table 2 and Fig. 7) comprise up to 11 times higher proportions of the assemblages at Stas F and A than at Cl and C2 (overall mean 3 times). Practically all these radiolarians are warm-water taxa, including well known indicators of
Radiolarian
vertical
distribution
211
patterns
Table 4. Variations in radiolarian relative abundances in the O-300 m layer of the four stations investigated. Body of the table indicates significant differences at P < 0.05 (one-way ANOVA and multiple contrasts according to Scheffi). Right-hand columns give the corresponding average percentages at each station (O-300 m only)
[14.77(300)1112.40 14 1Rhizoplegma 31
lheocatypha
boreale bicornk
0.52 (200)
0.34(200)
9.26(200)
0.65 (100)
WI “._. (200)
5.89
(100)
,I
5.89
1
p.12 500’ (
0.69 (0) 82 lLithe1iu.s sp. afi L. nautiloides
t t
135 45 24 54
16 21 117 20 89
165 134
1
Amphispyrti reticulota Potysolenia murrayana 1cornutelh profimda Heliodixus asteriscus ~Lamrocyclas maritalis $ichopilium bicorne 1Dictyophimus (?) sp. 1 Botryostrobus aquilonnrir 1Plagoniidae gen. et sp. indet. Lithopera bacca 1Callimitra cnro[ofae
0.: it 0.23 1 0.52 (lOa)
(4oo)l
1.83 (50)
~(1ooo)1.37(300)
11.72
11 (100)
(200)
IO.49 (300) 0.33 1.38 (0)
2mO1 1.37 (25)
1.38 (0)
0.52 (100)
91 1Siphocampe
--
98 1Phormostichoartus corbula 46 Bathropyramb woodringi 108 1Lithocampe sp. I
182 1Sethoconus
148 Lithe&u
(?) sp.
sp. 2
1 0.46 (50)
.-c-
0.
It- 0.06 (1000)
IO.09 ‘yoo’
1
1
M 1.38 1.37 0.77
arachnea
105 ~C)~tolagenalaguncu[a
1.72 1.55 1.38
212
S. A. KLING and D. BOLTOVSKOY
m
Station No
F (12
preference
spp.) (20
spp.)
0
c2 Stations
Cl
Fig. 11. Mean species-specific percentage contributions of the site-sensitive radiolarians to the G300 m assemblages of the four stations surveyed. Radiolarians are grouped according to their relative abundances at the four sites (cf. Table 4).
tropical-subtropical waters such as Didymocyrtis tetrathalamus, Eucyrtidium acuminaturn, Lipmanella dictyoceras, Pterocanium praetextum, Pterocanium trilobum, Pterocorys hertwigii, Sethophormis aurelia, Tetrapyle octacantha and Theocorythium trachelium. Although vertical profiles of these 12 radiolarians at A and F are very similar (Table 2)) a noteworthy difference is that their decrease with depth is much sharper at the coastal site (A), than at the oceanic one (F), in accordance with the difference in mixed-layer depths noted above. Thus, although at 0 m they comprise 32% (F) to 36% (A) of the fauna (unidentifieds excluded), at 50 m the percentage at F is 35%) and only 12% at A. In other words, similarities between F and A are high in the uppermost 25 m, but decrease rapidly with depth. This conclusion is clearly illustrated by results shown in Fig. 12. At 0 m Stas F and A are the most similar in their overall (relative) specific makeups. Below the surface, however, the oceanic Sta. F differs strongly from the other sites, and especially form Sta. A (Table 4). Some warm-water radiolarians, however, are abundant at the surface at Sta. F but show no substantial peaks at Sta. A. Among these, Cannobotryidae spp. et gen. indet., Spongosphaera streptacantha, Stylodictya spp. and Theopilium tricostatum, seem to be the most closely tied to site F; they account for 3.7% of total fauna at O-100 m at Sta. F, and only 0.6% at A. Pterocorys clausus and Theocorys veneris, also very abundant at the surface of Sta. F, have only very mild peaks at the surface of Sta. A. It is conceivable that all the above taxa are closely restricted to the Central Water and therefore cannot survive the trip to or conditions at Sta. A. Although the above interpretation is perhaps the most plausible, alternative hypotheses cannot be ruled out offhand. For example, the high abundances at Sta. F could represent a productivity event whose effects were not yet felt at Sta. A at the time of sampling. Comparison of our data with the pattern of polycystine distribution along the California Current derived by BOLTOVSKOY and RIEDEL (1987) reinforce the above conclusions. BOLTOVSKOY and RIEDEL (op. cit.), in a study of radiolarians from plankton tows collected
Radiolarian
vertical
distribution
patterns
213
Fig. 12. Similarities between overall (percentage) specific makeups at the four stations surveyed at 0, 50, 100 and 300 m. Clusters are based on MORISITA'S (1959) similarity index (HORN'S, 1966, modification), and UPGMA (SNEATH and SOKAL, 1973).
I
0.5
I
0.6
Fig. 13. Similarities between numbers for BOLTOVSKOY and California Current collected MORISITA'S (1959) similarity
I
/
1
I
0.7
0.8
0.9
1.0
radiolarian composition at 0 m at the stations surveyed and pooled RIEDEL'S (1987) station groups in O-LOO m plankton tows in the throughout 1972 (CC: California Current). Cluster is based on index (HORN'S, 1966, modification), and UPGMA (SNEATH and SOKAL, 1973).
throughout 1972 along the southern California Current grouped the assemblages retrieved into three distinct categories: colder-water associations restricted to the area north of 3234”N; transitional associations occurring between 25”N and 32-34”N; and warmer-water associations south of 25”N. Clustering of the above data (averaged numbers for pooled stations within each of the three categories), with 0 m results for our Stas (Fig. 13) shows
214
S. A.KLINC~~~D.BOLTOVSKOY
that sites F and A compare closely with the planktonic California Current warmer-water assemblage (“CC Warm” in Fig. 13), while Stas Cl and C2 cluster with the transitional assemblage of BOLTOVSKOYand RIEDEL (“CC Trans.“), and are also generally comparable to the colder-water one (“CC Cold”). The cold-water signal at subsurface layers of Sta. A may be reinforced by coastal upwelling, thus further enhancing the abundances of deeper-water radiolarians at this site. Such a mechanism is supported by the fact that, of the 11 radiolarians listed as “deep” (see “The O-2000 m layer” above), only two are present above 300 m at Sta. F (Spongopyle osculosa and Spongurus sp.), while at Sta. A nine of them occur in the same depthinterval. Similarly, some radiolarians that peak at subsurface depths at Stas F, C2 and Cl rise to a surface maximum at Sta. A (e.g. Dictyocoryneprofunda, Lithostrobus hexagonalis, and a few others, see Appendices 1,2). The inferred circulation responsible for radiolarian distribution patterns defined is schematically illustrated in Fig. 14. Implications
for the sedimentary record
Most distributional studies of Radiolaria based on surface sediment samples at some point dwell on comparisons between the bottom biogeographic imprint and hydrographic settings in the overlying mixed layer. These interpretations rely, either implicitly or explicitly, on the assumption that near-surface waters supply most or all of the shells recorded in bottom samples. Although the few direct comparisons of plankton and sedimentary information found important dissimilarities (PETRUSHEVSKAYA, 1971; RENZ, 1976; BOLTOVSKOY,1988, 1992; WELLING, 1990; SWANBERGand BJ~RKLUND, 1992),the latter were usually attributed to selective dissolution (PETRUSHEVSKAYA, 1981; RENZ, 1976; WELLING, 1990). However, a potentially important source of bias, or at least of blurring of differences between hydrographically dissimilar areas at the surface, is the flux of radiolarian shells from intermediate and deep layers. Because the physical environment in the meso- and bathypelagic realms is homogeneous throughout large areas of the World Ocean, its fauna is monotonous and it does not match biogeographic divisions in the upper waters. We attempted to compare the integrated absolute abundances of our shallow species, which are typically abundant over short depth intervals, with those of the deep radiolarians, which usually are less abundant over larger vertical ranges. Figure 15 illustrates the results of this analysis for Stas Cl and C2. Upper 50 m species (cf. Table 2) were assumed to yield live individuals between 0 and 100 m, while for the deep taxa (cf. Table 2) we averaged all their (presumably live) records between 300 and 2000 m. Although admittedly based on speculative assumptions, this figure suggests that depth-integrated abundances of deep-living radiolarians are higher than those of the shallow-water forms (approx. 4 times, according to data in Fig. 15). The integrated abundances are higher for the deep taxa (approx. 4 times at Sta. Cl, and about even at C2) even if calculated over 1000 rather than 2000 m. However, the contribution to sediment buildup from these deeper species, whose distributional patterns are not closely coupled with near-surface conditions, is dependent not only on their standing stocks, but also on their reproduction rates. It is conceivable that the nutrient-limiting conditions of meso- and bathypelagic depths render these sparse populations less productive than near-surface species. The degree to which these
Radiolarian
Fig.
14.
Inferred
circulation
vertical
affecting
distribution
the radiolarian
215
patterns
distribution
patterns
defined.
Radiolarian
cy 1.2 E 0-e nb 0.4 2 0.0
! ; r” % ‘pp~~qJ~ao:“‘““” r l-f-rrr
vertical
distribution
rrt-
0 ~ 0I- I-
Upper 50 m species at O-l 00 m Fig.
15.
217
patterns
N
Xr;E!
F 1?-
Deep species at 300-2000 m
Comparison of depth-integrated abundances of upper 50 m species, vs those of the deep species (cf. Table 2), at Stas Cl and C2. See Appendix 1 for species ID numbers.
presumably slower reproduction rates outweigh their higher absolute abundances remains an open question. The abundances of deep-water populations may be further enhanced by shells transported meridionally at depth from lower-latitude areas. As discussed above, several of the deeper-water radiolarians of our collection are absent or very scarce in O-100 m California Current waters throughout the year, but they are dominant components in north Pacific planktonic and sedimentary assemblages: Botryostrobus aquilonaris (ROBERTSON, 1975), Dictyophimus (?) sp. 1 (SACHS, 1973), Echinomma delicatulum (ROBERTSON, 1975), Heterucantha dentata (transitional species according to NIGRINI, 1970; SACHS, 1973; ROBERTSON, 1975), Rhizoplegma boreale (BLUEFORD, 1983), Siphocampe arachnea (KRUGLIKOVA, 1969; BLUEFORD, 1983), Spongurus sp. (TAKAHASHI, 1987, as Tholospyra group), Theocalyptra davisiana s.1. (cf. SACHS, 1973; BLUEFORD, 1983). These radiolarians seem to be abundant surface to subsurface dwellers closer to the pole but as the expatriated fractions of their populations move southward they sink deeper in the water-column and their densities decrease. This type of distribution pattern was demonstrated for some coldwater phaeodarian Radiolaria which submerge southward with the Intermediate subarctic waters (KLING, 1976), as well as for other radiolarians and zooplankton in general in all three oceans (VINOGRADOV, 1968; BOLTOVSKOY, 1992). BOLTOVSKOY (1988) reviewed mechanisms that can substantiate and enhance equatorward subsurface extensions of the ranges of cold-water planktonic organisms. An interesting example is Lithomitra arachnea, which is restricted to depths in excess of 200 m in our materials, peaking at 300-500 m (Sta. C2) to 1000 m (Stas Cl, F); and to lOO300 m in the central north Pacific (KLING, 1979). This species represents 25-60% of all radiolarians in surface sediments of the north Pacific north of 45-50”N, dropping to O-5% at the equator (KRUGLIKOVA, 1969). In sediments of the California Borderland it dominates the thanatocoenoses with contributions around 15-20% (KRUGLIKOVA, 1969; KLING, 1977). Yet in the O-100 m plankton it is absent altogether (BOLTOVSKOY and RIEDEL, 1987). The depths at which L. arachnea is found in extrapolar areas have low overall radiolarian standing stocks, and feeding conditions probably do not support very active growth and reproduction. Thus, we suggest that high abundances of this species in middle and low
218
S. A. KLING~~~
D.BOLTOVSKOY
latitude sediments are largely due to meridional transport, rather than to in situ production. Although vertical distribution profiles for L. aruchnea are not available for polar waters, the fact that it dominates high-latitude sedimentary assemblages implies high standing stocks of living populations which, therefore, are most probably associated with levels of maximum radiolarian abundance. In Arctic and Antarctic seas these maxima are located deeper than elsewhere, at about 20&400 m (TIBBS, 1967; BOLTOVSKOY and ALDER, 1992a,b; ALDER and BOLTOVSKOY, 1993). As mentioned in the introduction, the most widely used method in paleoenvironmental interpretations based on microfossil remains-the IMBRIE and KIPP (1971) factor analysis, transfer function technique-relies on the assumption that “core top faunas are systematically related to the physical nature of the overlying surface waters”. In this context, “systematically” means that the relationship in question has not varied in the time-span under investigation. However, the above-discussed contribution of laterally transported shells is dependent on the strength of intermediate and deep currents carrying-toward the equator-cooled waters that sink at the poles. Intensity of formation and output of these currents varied greatly between glacials and interglacials (BOYLE and KEIGWIN, 1987), conceivably affecting the contribution of cold-water polar shells to lower-latitude areas. Since meridional intermediate- and deep-water transport is strongest in the Atlantic and weakest in the Pacific, relationships between the sediments and the upper surface layer have probably been less uniform through time in the former than in the latter. Bathropyramis woodringi, Cornutella profunda, Cyrtolagena laguncula, and a few others, on the other hand, seem to be deep-water inhabitants in both low- and highlatitude areas. Their percentage contributions to overall sedimentary radiolarian assemblages do not increase towards the poles (NIGRINI, 1967; JOHNSON and NIGRINI, 1980,1982; BOLTOVSKOY, 1987). As far as paleoenvironmental interpretations are concerned, their records seem to be neutral background noise with no known information attached, other than the existence of abyssal ocean depths. REFERENCES V. A. and D. BOLTOVSKOY (1993) The ecology of larger microzooplankton in the Weddell-Scotia Confluence Area: horizontal and vertical distribution patterns. Journal of Marine l+arch, 51,323-344. BAKER A. DEC., M. R. CLARKE and M. J. HARRIS (1973) The N.I.O. combination net (RMT 1 + 8) and further developments of rectangular midwater trawls. Journal of the Marine Biological Association of the United Kingdom, 53, 167-184. BERNSTEIN T. (1934) Zooplankton Karskogo Morya po materialam ekspeditsii Arkticheskogo Instituta na “Sedove” 1930 goda i “Lomonosove” 1931 goda (Zooplankton des ntjrdlichen Teiles des Darischen Meeres). Trudy Arkticheskogo Instituta, Leningrad, 9, 3-58. BLUEFORD J. R. (1983) Distribution of Quaternary Radiolaria in the Navarin Basin geologic province, Bering Sea. Deep-Sea Research, 30,763-781. BOLTOVSKOY D. (1987) Sedimentary record of radiolarian biogeography in the equatorial to Antarctic western Pacific Ocean. Micropaleontology, 33,230-245. BOL~OVSKOY D. (1988) Equatorward sedimentary shadows of near-surface oceanographic patterns. Speculations in Science and Technology, 11,219-232. BOL~OVSKOY D. (1989) Las zonas de transici6n en la pelagial: biogeografia y paleobiogeografia. Mem. II Encontro Brasileiro de Plancton, Caiobd, 5-9 de dezembro de 1988, F. BRANDINI, editor, pp. 9-24. BOLTOVSKOY D. (1991) Holocene-upper Pleistocene radiolarian biogeography and paleoecology of the equatorial Pacific. Palaeogeography, Palaeoclimatology, Palaeoecology, 86,227-241. BOLTOVSKOY D. (1992) Current and productivity patterns in the equatorial Pacific across the Last Glacial Maximum based on radiolarian east-west and downcore fauna1 gradient. Micropaleontology, 38,397413. ALDER
Radiolarian
vertical
distribution
patterns
219
D. and V. A. ALDER (1992a) Paleoecological implications of radiolarian distribution and standing versus accumulation rates in the Weddell Sea. In: The Antarcticpaleoenvironment: a perspective on change, Antarctic Research Series, American Geophysical Union, 56, 377-384. BOLTOVSKOY D. and V. A. ALDER (1992b) Microzooplankton and tintinnid species-specific assemblage structures: patterns of distribution and year-to-year variations in the Weddell Sea (Antarctica). Journal of Plankton Research, 14, 1405-1423. BOLTOVSKOY D., V. A. ALDER and A. ABELMANN (1993a) Annual flux of radiolaria and other shelled plankters in the eastern equatorial Atlantic at 853 m: seasonal variations and polycystine species-specific responses. Deep-Sea Research I, 40, 1863-1895. BOLTOVSKOY D., V. A. ALDER and A. ABELMANN (1993b) Radiolarian sedimentary imprint in Atlantic equatorial sediments: comparison with the yearly flux at 853 m. Marine Micropaleontology, 23, l-12. BOLTOVSKOY D. and W. R. RIEDEL (1980) Polycystine Radiolaria from the Southwestern Atlantic Ocean plankton. Rev&a Espafiola de Micropaleontologia, 12,99-146. BOLTOVSKOY D. and W. R. RIEDEL (1987) Polycystine Radiolaria from the California Current region: seasonal and geographic patterns. Marine Micropaleontology, 12,655104. BOLTOVSKOY E. and H. A. LENA (1970) On the decomposition of the protoplasm and the sinking velocity of the planktonic foraminifers. Intationale Revue des Gesamten Hydrobiologie, 55,797-804. BOYLE E. A. and L. KEIGWIN (1987) North Atlantic thermohaline circulation during the past 20,000 years linked to high-latitude surface temperature. Nature, 330, 3540. BURCKLE L. H. (1981) Displaced Antarctic diatoms in the Almirante Passage. Marine Geology, 39, M39-M43. CARON D. A. and N. R. SWANBERG (1990) The ecology of planktonic sarcodines. Aquatic Science, 3, 147-180. CASEY R. E. (1966) A seasonal study on the distribution of polycystine radiolarians from waters overlying the Catalina Basin, Southern California. Ph.D. Thesis, University of Southern California, 136 pp. DWORETZKY B. A. and J. J. MORLEY (1987) Vertical distribution of Radiolaria in the eastern equatorial Atlantic: analysis of a multiple series of closely-spaced plankton tows. Marine Micropaleontology, 12, l-19. GOLL R. M. (1968) Classification and phylogeny of Trissocyclidae (Radiolaria) in the Pacific and Caribbean Basins. Journal of Paleontology, 42, 1409-1432. GOLL R. M. (1976) Morphological intergradation between Lophospyris and Phormospyris (Trissocyclidae, Radiolaria). Micropaleontology; 22, 379418. GOWING M. M. and S. L. COALE (1989) Fluxes of living radiolarians and their skeletons along a northeast Pacific transect from coastal upwelling to open ocean waters. Deep-Sea Research, 36,561-576. GOWING M. M. and D. L. GARRISON (1992) Abundance and feeding ecology of larger protozooplankton in the ice edge zone of the Weddell and Scotia Seas during the austral winter. Deep-Sea Research, 39,893-920. HAECKEL E. (1887) Report on the Radiolaria collected by H.M.S. Challenger during the years 1873-1876. Report on the Scientific Results of the Voyage of H.M.S. Challenger during the years 1873-1876, Zoology, 18, pp. i-clxxxviii + I-1803. HORN H. S. (1966) Measurement of “overlap” m comparative ecological studies. American Naturalist, 100, 419-424. IMBRIE J. and N. G. KIPP (1971) A new micropaleontological method for quantitative paleoclimatology: applications to a late Pleistocene Caribbean core. The Late Cenozoic Ages, Yale University Press, New Haven, pp. 71-181. JOHNSON D. A. and C. NIGRINI (1980) Radiolarian biogeography in surface sediments of the western Indian ocean. Marine Micropaleontology, 5, 111-152. JOHNSON D. A. and C. NIGRINI (1982) Radiolarian biogeography in surface sediments of the eastern Indian ocean. Marine Micropaleontology, ‘7, 237-281. JORGENSEN E. (1905) The protist plankton and the diatoms in bottom samples. Bergens Museum Skrifter, 1905, pp. 49-151, 195-225. KLING S. A. (1973) Radiolaria from the eastern north Pacific, Deep Sea Drilling Project, Leg 18. Initial Reports of the Deep-Sea Drilling Project, U.S. Government Printing Office, Washington, D.C., 18, 617-671. KLING S. A. (1976) Relation of radiolarian distribution to subsurface hydrography in the North Pacific. Deep-&a Research, 23, 1043-1058. KLING S. A. (1977) Local and regional imprints on radiolarian assemblages from California coastal basin sediments. Marine Micropaleontology, 2, 207-221. KLING S. A. (1979) Vertical distribution of polycystine radiolarians in the central North Pacific. Marine Micropaleontology, 4, 295-318. KRUGLIKOVA S. B. (1969) Radiolyarii v poverkhnostnom sloe osadkov severnoi poloviny Tikhogo Gkeana. In: BOLTOVSKOY
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and D. BOLTOVSKOY
Tikhii Okean, Mikroflora i mikrofauna v sovremennykh osadkakh Takhogo Okeana. Akademiya Nauk SSSR, Nauka, Moskva, pp. 48-72. MCMILLEN K. J. and R. E. CASEY (1978) Distribution of living polycystine radiolarians in the Gulf of Mexico and Caribbean Sea, and comparison with the sedimentary record. Marine Micropaleontology, 3, 121-145. MILLXMAN J. D. and K. TAKAHASHI (in press). Carbonate and opal production and accumulation in the ocean. In: GlobalsurJicialgeoJEuxes: Modern to Glacial, T. M. USSELMAN, W. HAY and M. MEYBECK, editors, National Academy Press. MORISITA M. (1959) Measuring of interspecific similarity and association between communities. Memoirs of the Faculty of Science, Kyushu University, Series E (Biology), 3,65-80. MORLEY J. J. and J. C. STEPIEN (1984) Siliceous microfauna in waters beneath Antarctic sea ice. Marine Ecology Progress Series, 19, 207-210. NICRINI C. A. (1967) Radiolaria in pelagic sediments from the Indian and Atlantic Oceans. Bulletin of the Scripps Institution of Oceanography, University of California, San Diego, 11, 125 pp. NIGRINI C. A. (1970) Radiolarian assemblages in the north Pacific and their application to a study of Quaternary sediments in core V20-130. Geological Society of America, Memoir 126, pp. 139-183. NIGRINI C. A. and T. C. MOORE (1979) A guide to Modern Radiolaria. Cushman Foundation for Foraminiferal Research, Special Publication 16, pp. Sl-S142 + Nl-N106. PELAEZ J. and J. A. MCGOWAN (1986) Phytoplankton pigment patterns in the California Current as determined by satellite. Limnology and Oceanography, 31,927-950. PETRUSHEVSKAYA M. G. (1967) Radiolyarii otryadov Spumellaria i Nassellaria Antarkticheskoi oblasti (po materiaiam Sovetskoi Antarkticheskoi Ekspeditzii). Issledovaniya Fauny Morei ZX(XII), Rezultaty Biologicheskikh Issledovanii Sovetskikh Antarkticheskikh Ekspeditsii 19.55-1958,3,5-186. PETRUSHEVSKAYA M. G. (1971) Spumellarian and nasselarian Radiolaria in the plankton and bottom sediments of the Central Pacific. In: The micropaleontology of oceans, B. M. FUNNELL and W. R. RIEDEL, editors, Cambridge University Press, Cambridge, pp. 309-318. PIELOU E. C. (1966) Shannon’s formula as a measure of species diversity: it’s use and misuse. American Naturalist, 100,463-465. POKRAS E. M. and B. MOLFINO (1986) Oceanographic control of diatom abundances and species distributions in surface sediments of the tropical and southeast Atlantic. Marine Micropaleontology, 10, 165-188. POPOFSKY A. (1908) Die Radiolarien des Antarktis (mit Ausnahme der Tripyleen). Deutsche Siidpolar Expedition 1901-1903,lO (Zool. 2), 3,183-305. POPOFSKY A. (1913) Die Nassellarien des Warmassergebietes. Deutsche Siidpolar Expedion 1901-1903,14 (Zool. 6) 217-416. REID J. L. (1965) Intermediate waters of the Pacific Ocean. The Johns Hopkins University Press, Baltimore, 85 PP. REID J. L. (1973) The shallow salinity minima of the Pacific Ocean. Deep-Sea Research, 20,51-68. REID J. L. G. I. RODEN and J. G. WYLLIE (1958) Studies of the California Current system. CalCOFI Progress Reports 7-l-56 to l-l-58, Marine Research Committee, California Department of Fish and Game, Sacramento, California, pp. 27-56. RENZ G. W. (1976) The distribution and ecology of Radiolaria in the Central Pacific; plankton and surface sediments. Bulletin of the Scripps Institution of Oceanography, 22, l-267. ROBERTSON J. H. (1975) Glacial to interglacial oceanographic changes in the northwest Pacific, including a continuous record of the last 400,000 years. Ph.D. Thesis, Columbia University, 355 pp. SACHS H. M. (1973) Quantitative radiolarian-based paleo-oceanography in late Pleistocene subarctic Pacific sediments. Ph.D. Thesis, Brown University, 201 pp. SHANNON C. E. and W. WEAVER (1949) Th e mathematical theory of communication. University of Illinois Press, Urbana, 125 pp. SNEATH P. H. A. and R. R. SOKAL (1973) Numerical taxonomy. Freeman, San Francisco, 573 pp. SWANBERG N. R. and K. R. BJ~RKLUND (1992) The radiolarian fauna of western Norwegian fjords: a multivariate comparison of the sediment and plankton assemblages. Micropaleontology, 38,57-74. TAKAHASHI K. (1983/84) Radiolaria: sinking population, standing stock, and production rate. Marine Micropaleontology, 8, 171-181. TAKAHASHI K. (1987) Radiolarian flux and seasonality: climatic and El Nido response in the Subarctic Pacific, 1982-1984. Global Biogeochemical Cycles, 1,213-231. TAKAHASHI K. and S. HONJO (1983) Radiolarian skeletons: size, weight, sinking speed, and residence time in tropical pelagic oceans. Deep-Sea Research, 30,543-568.
Radiolarian TIBBS J. F. (1967)
On some planktonic
protozoa
vertical taken
distribution from
the track
221
patterns of Drift
Station
ARLIS
I, 1960-1961.
Arctic
Institute of North America, 20, 247-254. VINOGRADOV M. E. (1968) Vertikalnoe raspredelenie okeanicheskogo zooplanktona. Nauka, L. A. (1990) Radiolarian microfauna in the northern California variability and implications for paleoceanographic reconstructions.
WELLING
Moskva, 320 pp. Current System: spatial and temporal M.Sc. Thesis, Oregon State University,
80 PP. J. G. (1966) Geostrophic Cooperative Oceanic Fisheries
WYLLIE
flow of the Investigations,
California Current at the surface and Atlas No 4, 13 pp, and 288 charts.
at 200 m. California
222
andD.
S. A. KLINC
APPENDIX
1.
BOLTOVSKOY
RADIOLARIAN
SPECIES
LIST
Species arc listed in alphabetical order, preceded by ID number. First set of square brackets gives maximum abundance recorded at each station (in ind. md3) and corresponding depth. Second set of square brackets furnishes a bibliographic identification reference and/or comments on the species’ taxonomic position. 91
Acunthodesmia sp. aff, A. vincufata (Mueller, Boltovskoy
and Riedel,
Giraffospyris
1987, Goll,
circumflsca,
41
Acunthodesmia
47
Acrosphuem spinosa (Haeckel,
1857)
as A. vinculutu
[F: 2.8-O;
Mueller.
C2: 0.5-50;
Open,
angular
Cl:
0.3-25;
structure,
A: 1.4-O]
similar
[?
to that of
19681
vincuhztu (Mueller,
1857)
[F: 1.4-O;
C2: ---; Cl:
---; A: 0.5-501
[Boltovskoy
and Riedel,
19871 Riedel, 181
Actinomma anturcticum (Haeckel, Rkdel,
68
132
[F: 0.1-750;
C2: 0.1-750;
Cl:
0.3-25;
A: ‘0.3-1001
C2: 3.7-50;
Cl:
20.0-o;
A: 2.8-O]
[Boltovskoy
and
1860)
[F: 5.5-O;
[Boltovskoy
and
19801
Actinomma leptodenum Echinomma
15
1860)
19801
(Joergensen,
leptodenmun
1900)
Joergensen;
[F: 0.5-50;
Boltovskoy
C2: 0.3-100;
and Riedel,
Cl:
---; A: ---I
[Kling,
1977,
as
19871
Actinomma sol Cleve, 1900 [F: 1.0-100; C2: ---; Cl: 0.5-50; A: 1.4-O] [Boltovskoy and Riedel, Actinommidae sp. et gen. indet. [F: ---; C2: ---; Cl: 0.1-1000; A: ---] [Various unidentified
19801
actinommids] 135
Amphiplectu acrostoma Haeckel, 1887 [F: ---; C2: 0. I-500; Cl: ---; A: ---1 [Petrushevskaya, Amphispyris reticuhzta (Ehrenberg, 1872) [F: 1.0-100; C2: 0.3-25; Cl: 0.2-500; A: 2.8-O]
77
Amphirhopdum
141
and Riedel,
19711 [Boltovskoy
19871
ypsilon Haeckel,
1887 [F:
1.4-O;
C2: ---; Cl:
0.2-200;
A: 1.4-O]
[Boltovskoy
and Riedel,
19871 170
Anthocyrtidium
40
Anthocyrtidium
1979;
84
Boltovskoy
same as specimens
somewhat
more
Amchnocomllium
with
C2: 0.3-25;
[F: 0.7-O;
Cl:
0.1-100;
a hyaline Cl:
0.2-50;
C2: ---; Cl:
0.1-400;
1908, except distally
coarser
Cl:
1987,
A: 0.2-3001 5.0-50;
under
A: 2.3-1001
the same name,
but
A: 2.3-501
[Probably
bears
same as Lithomelissa
to Psilomelissa
a slender
apical
spine,
tricuspidata
var.
and pores
on the
and irregular] Cl:
---; A: ---I [Similar
to but narrower
than
19711
1862 [F: 0.5-100;
C2: 1.4-50;
Cl:
0.3-25;
A: 0.3-1001
[Boltovskoy
19871
Amchnocorys (?) sp. aff. A. pentacantha Popofsky,
1913 [F: 4.8-O;
C2: 5.0-50;
Cl:
0.5-50;
19771
Artostrobus annul&us [Petrushevskaya,
[Kling,
peristome]
A: ---I [Similar
that the cephalis
sp. 2 [F: ---; C2: 0.2-300;
sp. in Pterushevskaya,
C2: 0.1-750;
and Riedel,
sp. 3 [F: 0.7-O;
Amchnoctwys circumtexta Haeckel,
[Kling, 185
1872)
19871
(?)
C2: 0.5-50;
by Boltovskoy
and Riedel,
become
Amchnocorullium
and Riedel, 169
illustrated
less campanulate,
of Popofsky,
thorax
11.7-O;
in Boltovskoy
Lophophaenoma 150
[F:
19871
sp. 1 [F: 2.8-O;
Amchnocomllium longer
ovate,
Haeckel
abdominalis 142
and Riedel,
1872)
zunguebaricum (Ehrenberg,
[Probably
thorucites 13
ophirense (Ehrenberg,
(Bailey, 19671
1856)
[F: 0.0-1000;
C2: 0.3-100;
Cl:
0.3-1ooO;
A: 0.2-3001
A: 1.4-O]
Radiolarian
46
Barhropyramis
20
[Kling, 19731 Borryosrrobus aquilonaris
woodringi
vertical
(Campbell
distribution
and Clark,
(Bailey,
1856)
1944)
223
patterns
[F: ---; C2: 0.5300;
[F: 0.1-1000;
C2: 1.4-300;
Cl:
Cl:
0.6-400;
1.7-200;
A: 0.5-3001
A: 0.5-3001
[Kling,
19791 9.5
Botryostrobus uurituduustrulis [Kling,
1979,
(Ehrenberg,
as B. au&us-ausrralis
carolotae Haeckel,
134
&&r&m
1887 [F:
17
? Culoeyclas monumentum Haeckel,
18
Cunnobotlyidae
1844)
[F: 6.2-100;
group;
Boltovskoy
1.4-25;
C2: 0.1-400;
C2:
15.5-50;
and Riedel, Cl:
Cl:
1987;
11.7-50;
A: 12.8-1001
Boltovskoy
---; A: 1.4-O]
and Vrba,
[Boltovskoy
19891
and Riedel,
19871 spp.
1887 [F: 0.5-100;
et gen. indet.
[F:
14.5-O;
C2: 0.3-100;
C2: 0.3-25;
Cl:
Cl:
2.1-50;
---; A: ---I [Haeckel,
18871
A: 0.5-501
[Various
unidentified
to Carpocanistrum
spp.
A and B in
catmobotryids] 119
Carpocanium sp. 1 [F: 4.1-O; Boltovskoy
118
and Riedel,
Curpocanium sp. 2 [F: 6.9-O; and B in Boltovskoy
4
C2: 0.9-50;
Cl:
0.9-200;
A: 1.6-1001
1987; with
tennina
teeth]
(Popofsky,
1913)
[Similar
[F: ---; C2: 0.1-500;
Carposphuem sp. aff. C. acaathophom (Popofsky, [Similar
to C. acanthophora
megaceros Hollande
Cladococcus [Boltovskoy
87
A: ---I [Similar
Cl:
to Carpocanistrum
0.1-400;
A: ---I
spp.
A
[Boltovskoy
and
19871
0.3-3001 56
2.4-25;
no teeth]
and Riedel,
Curposphaem ncunthophom Riedel,
60
C2: ---; Cl:
1987;
and Riedel,
Clathrocunium Riedel,
1980;
1913)
in Boltovskoy
[F: 0.7-O;
and Enjumet,
1960 [F: 4.6-50;
Kling,
as Cladococcus
courctutum Ehrenberg,
1977, 1860 [F:
C2: 0.2-300;
and Riedel,
8.3-O;
1987,
Cl:
but with
C2: 4.150;
1.2-200;
larger
Cl:
A:
pores]
1.7-200;
A: 4.1-5011
spp.]
C2: 1.0-25;
Cl:
0.7-O;
A: 12.4-O]
[Boltovskoy
and
19871
160
Collosphaem macropom Popofsky,
24
Cornutellaprofunda
1917 [F: 0.2-200;
C2: ---; Cl:
---; A: 8.3-O]
[Boltovskoy
and Riedel,
19871 Boltovskoy 151
123
[Somewhat
more
C. cervus;
includes
Corocdyptru
sp. aff.
83
cyrtohzgena 1979,
33
1873,
C. danaes
and Riedel,
group
[F:
than the category
Eucecryphalus
Kling,
Cl:
2.7-200;
A: 0.2-3001
[Kling,
1979;
craspedota Haeckel,
1.0-200;
illustrated
C2: 16.3-100;
Cl:
by Boltovskoy
(Joergensen)
1887 [F: 0.5-100;
22.4-50;
and Riedel,
of Kling,
A: 16.4-1001 1987,
under
the name
19791
C2: 0.3-300;
Cl:
0.7-25;
A: 2.7-501
19871 C2: 0.2-300;
Cl:
0.2-50;
A: 0.5-SO]
[Petmshevskaya,
19771
Cubotholus SP. [F: 0.1-1000; Cypussis irre&?daris Nigrini, 1977,
105
variable
C2: 0.5-300;
Cromyechinus borealis (Cleve, 1899) [F: 0.3-300; 1967;
71
1854 [F: 0.1-500;
19871
Corocalyptra cerws Ehrenberg,
[Boltovskoy 11
Ehrenberg,
and Riedel,
C2: ---; Cl: 1%8
0.1-750;
[F: 6.9-25;
A: ---I [Unidentified
C2: 0.3-25;
Cl:
Tholoniidae]
1.4-O;
A: 4.1-O)
Cl:
0.3-1000;
[Nigrini,
1968;
Klmg,
19791 S%Z~UIK.U~~ Haeckel,
as Stichopera
pecrinata
Dictyocephduspapi~losus
1887 [F: 0.1-1000; Haeckel
(Ehrenberg,
C2: 0.2-500;
[Kbg,
A: 0.3-1001
group] 1872)
[F:
1.0-300;
C2:
1.0-200;
Cl:
1.7-200;
A: 1.4-O]
[Nigini,
19671 79
Dictyocorynepmfbfda
102
Dictyophimus crisi~e Ehrenberg, Dictywhimus @wcif&es (Bailey,
Hymeniastrum 32
1977,
1979,
Ehrenberg,
euclidis;
1860 [F: 3.4-25;
Boltovskoy
as Psertdodictyophimus
and Riedel,
C2: 2.7-50;
[F: 25.9-100; gracilipes;
5.2-25;
A: 9.7-O]
[Kling,
1977,
19871
1854 [F: ---; C2: 0.1-750; 1856)
Cl:
Cl:
0.1-2000;
C2: 48.0-100; Boltovskoy
Cl:
and Riedel,
A: ---I 11.9-50; 19871
[Nig&i,
19671
A: 35.3-1001
[Khng,
as
224 146 103 8 117 130 55 69 10 131 104
152 78 36 171 139 136 110 54 43 48 59 1
66
S. A. KLING
and D. BOLTOVSKOY
Dic~ophimus infibticatus Nigrini, 1968 [F: 0.5-200; C2: 7.5-100; Cl: 1.0-200; A: 5.6-lOO] [Khng, 1977, 19791 Dictyophimus (?) kilhnati (Renz, 1976) [F: 0.9-300; C2: ---; Cl: 0.2-300; A: 0.7-lOO] [Renz, 1976, as Corocalyptra killmari] Dictyophbnus (?) sp. 3 [F: 0.7-O; C2: ---; Cl: 0.3-25; A: ---1 [Similar to Dictyophimus sp. aff. D. infabricatus Nigrini in Boltovskoy and Riedel, 19871 Dictyophimus (?) sp. 1 [F: 0.5-100; C2: ---; Cl: 1.7-500; A: ---I [Similar to Dictyophimus multispinus in Bernstein, 19341 Dictyophitnus (?) sp. 2 [F: 8.8-100; C2: 0.2-300; Cl: ---; A: ---] [Resembles Dictyophimus hirundo (Haeckel) group in Nigrini and Moore, 19791 Didymocyrtis tetmthnfamus (Haeckel, 1887) IF: 40.7-o; C2: 2.1-25; Cl: 18.5-25; A: 33. l-O] [Kling, 1979, as Ommatartus tetrathalamus; Boltovskoy and Riedel, 19871 Drynwsphaem (?) sp. [F: 0.7-O; C2: ---; Cl: ---; A: ---] [Provisional generic assignment according to Haeckel, 18871 Echinomma delicatulum (Dogiel, 1952) [F: ---; C2: 0.0-2000; Cl: 0.1-1000; A: ---I [Petrushevskaya, 1967; Khng, 19771 Echinommapopofskii Petrushevskaya, 1967 [F: 0.2-300; C2: 0.7-O; Cl: 1.4-O; A: 0.5-3001 [Khng, 19771 Enneaphonnis (?) sp. [F: 0.9-300; C2: 1.0-200; Cl: 0.7-O; A: 0.3-3001 [Generally similar to Enneaphonnis spp. in Petrushevskaya, 1971; probably conspecific with Sethophormis rot&a Haeckel in Renz, 19761 Eucecvphnlus sp. [F: 0.5-100; C2: 24.5-W; Cl: 30.6-50; A: 53.7-1001 [Boltovskoy and Riedel, 19871 Euchitonia eZeguns/furcuta (Ehrenberg, 1872) [F: 2.8-O; C2: ---; Cl: 0.7-25; A: 1.4-O] [Boltovskoy and Riedel, 19871 Eucyrtidium acuminatum (Ehrenberg, 1844) [F: 122.7-O; C2: 2.4-25; Cl: 8.6-25; A: 24.8-O] [Nigrini, 1967; Nigrini and Moore, 19791 Eucyrtidium hexagon&urn Haeckel, 1887 [F: 0.7-25; C2: ---; Cl: 0.325; A: ---I [Nigrini, 1967; Nigrini and Moore, 19791 EucyMium hexustichum (Haeckel, 1887) [F: 4.1-100; C2: 3.250; Cl: 3.1-25; A: 4.6-501 [Boltovskoy and Riedel, 19871 EucyrtiAum sp. [F: ---; C2: ---; Cl: 0.1-1000; A: ---I [Similar to E. acuminatum but narrower and with longer abdomen] Eucyrtidium sp. cf. E. culvertense Martin, 1904 [F: ---; C2: ---; Cl: ---; A: 0.2-3001 [Probably same as E. cienkowskii Haeckel in Boltovskoy and Riedel, 19801 Heliodiscus astetiscus Haeckel, 1887 [F: 2.1-O; C2: 0.1-400; Cl: 0.3-200; A: ---I [&Zing, 1979; Boltovskoy and Riedel, 19871 Helotholus histricosu Joergensen, 1905 [F: 2.1-O; C2: 0.9-50; Cl: 0.7-25; A: 0.7-1001 [Rling, 1977; Boltovskoy and Riedel, 19871 Hetemcantha dentutu Mast, 1910 [F: ---; C2: 0.1-750; Cl: ---; A: 0.3-lOO] [Nigrini, 1970; also similar to several other described taza, such as Cladococcuspinetum in Haeckel, 18871 Hexucontium laevigutum Haeckel, 1887 [F: 0.5-100; C2: 1.4-50; Cl: 0.3-1000; A: ---I [Boltovskoy and Riedel, 19871 Hexncontium sp. 1 [F: 4.1-100; C2: 6.2-300; Cl: 10.1-50; A: 5.5-501 [Kling, 1977, as H. pachydennum; large pores and conspicuous by-spines, similar to H. pachydennum in Joergensen, 1905, and to H. hostile Cleve in Boltovskoy and Riedel, 19801 Hexacontium sp. 2 [F: 0.5-300; C2: 2.3-50; Cl: 0.2-50; A: 0.5-501 [Probably same as H. entacanthum in Rling, 1977, and Boltovskoy and Riedel, 1980, 19871
Radiolarian
28
Lamprocyttis nigriniae (Caulet,
vertical
1971)
distribution
[F: 0.9-300;
225
patterns
C2: 2.3-100;
Cl:
3.3-200;
A: 1.3-3001
[Kbng,
1977,
19791 52 129
Lampromitmpambolica Popofsky, Lnmpromitm quadricuspis Haeckel, [Boltovskoy
16 72
and Riedel,
1913 [F: 0.5-100;
C2: 0.550;
1887 [F: 0.5-100;
C2:
Cl:
1.3-100;
---;
Cl:
A: 0.3-lOtI] 1.5-200;
[Popofsky,
19871
Lamprocycbzs maritalis Haeckel, 1887 [F: 0.3-1000; C2: 0.3-100; Cl: 2.1-200; A: ---I [Kling, L. m. polypora Nigrini] Larcopyfe buetschlii Dreyer, 1889 [F: 0.1-750; C2: 0.7-300; Cl: 9.3-200; A: 0.7-1001 [Kling, Boltovskoy
and Riedel
19131
A: 6.2-1001 1979,
as
1977;
19871
81
Larcopyle (2) sp. [F: ---; C2: 1.0-300; Cl: 0.5-300; A: 0.3-3001 [Conspicuously coarser external lattice than in Larcopyle buetschlii] Larcospim quadtanguta Haeckel, 1887 [F: 2.8-O; C2: 1.8-50; Cl: 6.2-25; A: 5.5-O] [Kling, 1977, 1979;
133
Lipmaneffa dictyocems (Haeckel,
9
Boltovskoy 1979;
and Riedel,
Boltovskoy
19871
and Riedel,
[F: 26.9-O;
as L. virchowii (Haeckel)]
C2: 9.1-50;
Cl:
7.6-25;
A: 40.0-O]
[Kling,
1977,
5
Lithamchnium
3
165
Lithelius minor Joergensen, 1900 [F: 2.1-25; C2: 1.6-100; Cl: 2.6-200; A: 13.8-O] [Kling, 1977, as L. minor (?)] Lithe&s sp. 1 [F: ---; C2: 0.2-300; Cl: ---; A: ---I [Unidentified Litheliidae, similar to Lithe&s sp. aff. L. alveolina Haeckel in Renz, 19761 Lithelius sp. 2 [F: ---; C2: 0.1-1000; Cl: 0.2-200; A: ---I [Unidentified Litheliidae; larger, coarser-meshed and shorter-spined than LitheZius sp. I] Lithelius sp. aff. L. nautiloides Popofsky, 1908 [F: 3.4-O; C2: 0.5-50; Cl: 0.4-1000; A: 1.4-O] [Probably conspecific with ?Pylospyra octopyle in Boltovskoy and Riedel, 19801 Lithocampe sp. 1 [F: 0.5-50; C2: ---; Cl: 0.1-1000; A: ---I [Nigrini, 1967, as Lithocumpe sp.] Lithomelissa setosa Joergensen, 1900 [F: 1.4-O; C2: 2.6-100; Cl: 11.7-25; A: 1.4-501 [Kling, 19771 Lithopem bacca Ehrenberg, 1872 [F: 0.7-O; C2: 0.1-750; Cl: 0.3-25; A: 1.4-O] [Kling, 1979; Boltovskoy
39
Lithostrobus haxagonalis Haeckel,
1979;
80 148 82 108 23
tentorium Haeckel,
1887) 1987,
Boltovskoy
and Riedel, Riedel, 154
1987;
and Riedel,
includes
C2: 5.2-100;
Cl:
7.1-50;
A: lO.l-1001
[Kling,
19871
many juvenile
specimens]
1887 [F: 0.5-100;
C2: 1.8-50;
1913 [F: 0.5-100;
C2: 0.2-300;
1887)
C2: 7.850;
Cl:
2.1-50;
A: 1.4-O]
[Boltovskoy
and
19871
Lophocoryspolyacantha Icling,
1860 [F: 3.1-100;
Popofsky,
Cl:
0.9-200;
A: ---I [Popofsky,
1913;
19791
168
Neosemantis distephanus (Haeckel,
92
Nephrospyris renifta Haeckel, 1887 [F: 1.4-O; C2: ---; Cl: ---; A: 0.3-1001 [Boltovskoy and Riedel, 19871 Porodiscus microporus (Stoehr, 1880) [F: 0.2-200; C2: 0.7-O; Cl: 1.4-O; A: 0.9-501 [Renq 19761 Peridium (?) SP. [F: 30.3-o; C2: 42.1-50; Cl: 49.1-25; A: 31.7-O] [Probably same as P. spinipes I-&&e1
Boltovskoy 74 85
in Casey, 70 184 38 50 98
and Riedel,
[F: 19.3-O;
Cl:
9.6-50;
A: 22.1-O]
[KJing,
1979;
19871
19661
Perypimmis circumtexta Haeckel, 1887 [F: Phormospytis s&bibs capoi Goll, 1976 [F: Phormospyris stabilis scuphipes GOB, 1976 Phormospyris stabilis stabifis (Goll, 1968) Phormostichoartus corbula (Harting, 1863)
---; C2: 0.2-300;
Cl:
---; C2: ---; Cl:
1.4-50;
[F: 0.5-100; [F: 2.1-O;
C2:
A: ---I
A: 3.2-501
1.4-50;
C2: 0.2-200;
[F: 0.1-1000;
0.2-300; Cl: Cl:
C2: 0.3-300;
2.5-50; 0.5-200; Cl:
19791
[Ming,
[GOB,
19761
A. 5.9-1001 A: 1.4-O]
0.6-1000;
[Go&
[KJQ,
19761 19791
A: 0.3~1001
[~hn~,
19791
89
Plagoniidae
gen. et sp. indet. [F: 1.6-100;
C2: 0.1-500;
Cl:
---; A: 0.3-1001
[Similar
to PI. 30, fig.
13 j,,
226 157 122 45 138
127 101 99 180 44 25 115 112 187 178 14 182
121 51 97 124 161 186 58 57
S.A. KLING andD. BOLTOVSKOY Popofsky, 19081 Plectauzntha (?) sp. aff. P. cremastoplegma Nigrini, 1968 [F: 17.9-O; C2: 0.2-300; Cl: 1.5-200; A: 5.5-O] [Provisional assignment according to Nigrini, 19681 Plegmosphaem sp. aff. P. lepticali Renz, 1976 [F: ---; C2: 0.9-300; Cl: 3.3-200; A: ---1 [Renz, 19761 Polysolenia murrayarm (Haeckel, 1887) [F: 2.7-25; C2: 0.0-1000; Cl: 0.1-500; A: 0.7-1001 [Boltovskoy and Riedel, 1987, as Acrosphaeru murrayuna] Psifomelissa (?) sp. [F: 19.3-O; C2: 7.8-50; Cl: 8.2-25; A: 2.8-O] [Probably related to Lophophaena sp. aff. L. apiculata Ehrenberg in Boltovskoy and Riedel, 1987; and to Psilomelissa galeata Ehrenberg in Popofsky, 19081 Pterocanium korotnevi (Dogiel, 1952) [F: 3.4-25; C2: 1.3-W; Cl: 2.1-O; A: 0.3-1001 [Nigrini and Moore, 19791 Pterocaniumpmetextum (Ehrenberg, 1872) [F: 36.4-25; C2: 0.5-50; Cl: 6.9-25; A: 40.0-O] [Boltovskoy and Riedel, 19871 Pterocanium sp. aff. P. gmndiporus Nigrini, 1968 [F: ---; C2: ---; Cl: 1.8-50; A: ---I [Similar to specimens illustrated by Nigrini, 1968, as P. gradiporus] Pterocanium sp. aff. P. trilobum (Haeckel, 1860) [F: 2.9-200; C2: ---; Cl: 1.025; A: 2.0-1001 [Atypical specimens resembling closely the named species] Pterocanium trilobum (Haeckel, 1860) [F: 37.2-O; C2: 2.4-25; Cl: 17.2-O; A: 26.2-O] [Kling, 19791 Pterocorys clausus (Popofsky, 1913) [F: 66.2-O; C2: 1.4-O; Cl: 2.1-O; A: 4.3-1001 [Klmg, 19791 Pterocorys hertwigii (Haeckel, 1887) [F: 41.4-O; C2: 4.1-25; Cl: 15.8-25; A: 63.5-O] [Boltovskoy and Riedel, 19871 Pterocorys minythomx (Nigrini, 1968) [F: 26.9-O; C2: 42.550; Cl: 50.1-25; A: 41.6-1001 [Kling, 1977, as Theoconus minythorax Nigrini; Boltovskoy and Riedel, 19871 Pterocorys sp. 1 [F: 1.8-50; C2: 6.5-25; Cl: 0.2-200; A: 1.6-1001 [Similar to P. macrocerus in Petrushevskaya, 19711 Pterocorys sp. 2 [F: 6.9-25; C2: 0.3-25; Cl: 0.3-25; A: 2.8-O] [Probably same as P. zancleus Mueller in Petrusbevskaya, 19711 Rhizoplegma boreale (Cleve, 1899) [F: 0.1-500; C2: ---; Cl: 0.2-200; A: 5.9-lOO] [Kling, 19771 Sethoconus (?) sp. [F: 0.1-1000; C2: 0.1-500; Cl: 0.3-1000; A: 0.3-1001 [Similar to Stichopilium variubile spinosum Popofsky in Petrushevskaya, 1967, but lacking postcephalic chamber divisions of Popofsky’s form] Sethoconus (?) sp. cf. S. (?) dogieli Petrushevskaya, 1967 [F: 0.1-500; C2: 0.2-200; Cl: 0.2-500; A: ---I. [Petrushevskaya, 19671 Sethophonnis aurelia Haeckel, 1887 [F: 71.7-O; C2: 3.7-50; Cl: 2.4-25; A: 46.9-O] [Boltovskoy and Riedel, 19871 Siphocampe arachnea (Ehrenberg, 1861) [F: 0.1-1000; C2: 0.2-300; Cl: 0.8-1000; A: ---I [Kling, 1979, as 5. aruchneu (Ehrenberg) group] Siphonosphaempolysiphonia Haeckel, 1887 [F: 3.4-25; C2: ---; Cl: 3.4-O; A: 5.5-O] [Kling, 1979; Boltovskoy and Riedel, 19871 Solenosphuempolymorpha (Haeckel, 1887) [F: 1.4-O; C2: ---; Cl: ---; A: ---I [Boltovskoy and Riedel, 19871 Spirocyrfis scalaris Haeckel, 1887 [F: 0.7-O; C2: 13.3-50; Cl: 5.3-50; A: 2.0-lOO] [Boltovskoy and Riedel, 1987, as S. scalariskomutella Haeckel (group ?)] Spongaster tetms Ehrenberg, 1860 irregdaris Nigrini, 1967 [F: 0.7-O; C2: 0.5-50; Cl: 4.1-25; A: 4.1-O] [Boltovskoy and Riedel, 19871 Spongocore cylitufriea Haeckel, 1860 [F: 8.8-100; C2: 1.4-O; Cl: 2.8-O; A: 7.8-501 [Kling, 1977, 1979,
Radiolarian
125
7
and Riedel,
227
patterns
C2: 8.6-300;
Cl:
8.9-25;
A: 4.3-1001
[Kling,
1979;
1987
Spongopyle osculosa Dreyer, 1889 [F: 3.0-400; C2: 2.4-300; Cl: 14.8-300; A: 12.4-3001 [Kling, 19771 Spongosphaera streptucantha Haeckel, 1862 [F: 11.7-O; C2: 2.7-50; Cl: 0.7-O; A: 0.9-501 [Boltovskoy and Riedel,
75
distribution
as S. puella; Boltovskoy and Riedel, 19871 Spongodiscus resurgens Ehrenberg, 1854 [F: 3.4-O; Boltovskoy
76
vertical
19871
Spongotrochus glacialis Popofsky,
1908 [F: 7.6-O;
C2: ---; Cl:
1.1-50;
A: 5.5-O]
[Boltovskoy
and Riedel,
19871 21
Sportgurus sp. [F: 1.5-500; C2: 2.9-500; Cl: 6.2-500; A: 5.5-O] [Kling, 1977, as Spongurus (?) sp.] Stichopilium bicorne Haeckel, 1887 [F: 0.2-400; C2: 1.8-50; Cl: 0.3-loo0, A: 0.3-1001 [Kling, 1979;
62
Stylacontatium bispiculum Popofsky,
27
Boltovskoy figs. 35
and Riedel,
19871 1912 [F: ---; C2: ---; Cl:
0.1-500;
A: ---I
[Skiing,
1973,
Plate 15,
1 l-141
Stylochlamydium venusturn Bailey, 1979;
Boltovskoy
and Riedel,
1856 [F: 0.9-50;
C2: 2.2-300;
Cl:
5.0-200;
A: 4.1-O]
[Wing,
1977,
19871
6
Stylodictya spp. [F: 11.0-o; C2: 2.2-200; Cl: 4.5-200; A: 3.9-1001 [Includes mainly forms similar to S. validispina in Kling, 19791 Tetrapyle octacantha Mueller, 1858 [F: 99.3-O; C2: 18.9-25; Cl: 37.8-25; A: 57.9-O] [Kling, 1979;
37
Theoculyptm bicornis (Popofsky,
73
Boltovskoy
and Riedel,
19871 1908)
[F: 0.5-200;
C2: 0.3-200;
Cl:
4.1-500;
A: 5.4-3003
[Kling,
1977,
19791 147
Theocalyptm aiwisiana (Ehrenberg, Cl:
106
29.9-200;
venen’s
Haeckel,
Boltovskoy
1967 [F: 0.9-300;
C2:
14.4-200;
19791 1862)
1887 [F: 41.4-O;
and Riedel,
[F: ---; C2: 0.7-200;
Cl:
0.5-100;
A: 2.9-1001
Boltovskoy
and Riedel
Tkeopilium tricostutum Haeckel, Riedel,
C2: 5.0-50;
Cl:
1.8-50;
A: 2.9-1001
[Rem,
1976;
W&g,
as Corocalyptra columba (Haeckel)]
1987,
Theocorythium tmchelium (Ehrenberg, 1979;
155
cornutoides Petrushevskaya,
1977,
19771
Theocovs 1979;
111
1862)
[Kling,
Theoculyptm davisiana davisiana (Ehrenberg, [Kling,
137
A: 3.1-3001
1872)
[F: 36.5-O;
C2: 0.5-50;
Cl:
4.8-25;
A: 15.2-O]
[a&
19871 1887 [F: 33.1-O;
C2: 25.6-50;
Cl:
2.1-200;
A: 2.3-501
[Boltovskoy
19871
90
zygocircusproductus
164
zygocircus sp. cf. 2. capulosus Popofsky,
and Riedel,
(Herhvig,
1879)
[F: 22.8-100;
C2: 1.4-50;
Cl:
1.5-200;
A: 5.9-1001
[Boltovskoy
19871
[Petrushevskaya,
1913 [F: 0.5-100;
C2: 0.2-300;
Cl:
0.2-200;
19711
163
Zygocircus (?) sp. [F: 4.1-O;
153
Unidentified
radiolarians
C2: 0.5-300; [F: 680.5-o;
Cl:
2.1-29;
C2: 241.2-100;
A: 1.4-O] Cl:
[D-shapedrings]
316.7-25;
A: 429.0-O]
A: 1.8-501
and
S.
A. KLING andD. BOLTOVSKOY APPENDIX
Radiolarian A 0
A 50
0.w 5.48 13.80 o.%? 0.00 0.00 57.94
0.00 0.w 0.00 0.00 0.00 0.00 1.38 1.38 0.043 0.04 0.W 0.00 0.00 0.00 0.00 2.78 5.52 0.04 1.38 1.38 4.14 24..%3 0.00 0.00 1.38 0.00 28.2, 0.W 0.03 0.00 1.38 0.00 0.00 33.1, 0.00
(ind. rnm3; top row gives station
counts
0.00 1.37
A loo 4.01 0.33 0.00 10.14
16.03 7.85 0.02 0.33 0.00 0.00 0.w 0.00 0.00 0.00 0.48 0.00 0.04 0.W 2.29 5.88 0.48 0.00 0.00 0.00 0.00 0.00 0.48 0.33 0.00 0.00 0.00 0.33 1.37 1.a, 0.00 0.W 4.25 0.W 0.33 0.02 0.00 0.04 .5.49 35.34 0.00 0.00 2.95 1.37 2.75 1.3, 0.00 0.33 3.66 5.80 0.02 0.65 0.02 2.29 0.40 0.W 0.00 0.65 1.37 7.85 0.W 0.65 0.00 0.04 0.00 0.33 0.00 0.33 0.00 2.29 0.00 0.04 3.86 4.12
A 300 2.28 3.43 0.00 2.12 4.24 0.16 0.w 0.33 0.00
0.16 0.00 0.33 0.16 0.00 0.00 0.00 0.40 0.00 0.10 0.10 0.00 3.10 1.31 4.4, 0.W 0.88 0.82 5.39 0.48 0.00 0.W 0.16 0.18 0.33 0.04 0.40 0.16 0.W
Cl 0 2.07 0.00 0.00 0.00 14.40 0.60 0.00 0.00 0.00
0.04 0.00 0.00 0.00 0.W 0.00 0.M) 0.00 0.00 2.07
0.W 2.07 1.38 0.00 4.83 0.00 4.83 2.78
0.00 0.00 0.04 0.00 0.00 0.00 17.24 0.00 0.W 0.00 0.00
7.78
4.14 0.00 0.00 0.04 0.00 0.00 4.14 0.w 0.00 0.W 5.52 4.14 1.38 1.38 %B8 0.09 5.52 1.38 0.W 1.38 31.73
-
0.48 0.00 0.00 0.00 0.48 0.00 0.00 0.00 0.00 O.MI 0.02 0.02 0.46 1.37
0.W 0.00 1.37
0.00 0.48 0.00 0.00 2.29 28.83
0.00 0.33 4.25 0.00 0.00 0.33 0.00 0.33 0.00 0.00 0.33 8.04
0.10 1.3 0.00 0.04 12.40 0.W 0.10 0.00 0.00 0.04 1.14 2.76 003 O.Oil 0.33 1.38 0.W 0.00 0.00 0.00 0.04 0.00 0.48 25.52
C-l 25
1.72
Cl 50
C-l loo
2
and depth, C-l 200
C-l 3co
left-hand C-l 400
column c-1 500
10.00 1.61 6.88 0.52 2.05 3.78 0.04 om 2.57 2.23 0.34 0.46 0.w 0.w 0.00 0.03 0.1, 0.00 7.09 0.6s 3.00 0.24 0.68 2.29 37.78 10.51 1.78 10.90 3.26 1.7, 1.37 0.34 0.23 0.00 0.17 0.17 0.23 0.23 0.34 0.00 0.08 0.17 0.00 0.w 0.00 0.w 0.00 0.00 0.34 0.52 0.00 0.11 0.00 o.cQ 0.00 0.00 0.00 0.00 0.00 0.17 0.00 0.23 0.1, 0.03 0.1, 0.1, 0.00 0.00 0.w 003 0.00 0.1, 0.w 0.00 0.00 0.00 0.17 0.02 0.00 0.00 0.00 0.46 0.53 0.17 0.00 0.00 0.00 0.00 0.23 0.00 2.08 0.00 0.00 0.23 0.00 0.W 0.00 0.03 0.00 0.00 0.00 0.69 2.06 0.00 0.17 0.17 0.M) 0.1, 0.6S 0.00 0.00 1.72 1.03 0.23 0.34 0.00 0.W 0.00 0.00 0.17 0.00 0.23 11.88 4.34 0.34 5.83 1.20 1.02 1.83 0.34 0.23 0.00 2.74 1.88 0.57 0.57 0.00 0.00 0.W 0.17 0.00 0.W 0.00 0.04 0.08 0.34 1.37 3.04 0.34 0.00 0.00 3.28 0.17 0.23 4.81 11.89 230 11.32 2.23 0.34 0.34 0.00 0.00 1.72 0.86 0.34 1.37 1.14 0.57 4.98 3.09 0.34 1.14 8.59 2.20 0.17 1.03 0.52 0.11 0.82 0.00 0.*3 0.03 1.03 0.00 0.57 4.12 0.34 2.51 0.17 0.5, 0.00 0.11 O.BS 0.00 2.08 0.W 0.88 0.W 0.23 0.46 0.00 5.03 0.48 0.00 0.00 0.00 0.03 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.89 0.00 0.08 0.5, 0.52 0.00 0.00 12.7, 10.29 1.08 2.57 0.00 0.1, 0.23 0.W O.Cil 0.06 0.00 0.00 0.00 0.11 0.00 0.00 0.00 0.17 0.34 0.57 0.34 0.34 0.00 0.M) 0.09 0.00 0.00 0.00 0.04 003 0.00 0.00 0.00 0.03 0.00 0.00 0.23 0.00 0.51 0.00 0.M) 0.W 2.40 080 0.1, 0.34 0.34 0.11 0.11 0.00 0.00 0.03 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.34 0.17 0.00 0.00 18.55 1.14 0.17 3.77 0.52 0.1, 0.11 0.29 1.72 0.34 0.23 0.48 0.68 0.9, 0.6S 2.06 0.1, 2.08 0.17 0.1, 0.48 4.12 0.46 003 0.89 0.34 0.00 0.00 0.W 0.23 0.09 0.00 0.17 0.23 0.23 0.00 0.00 0.00 1.20 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.1, 0.11 0.00 0.23 0.00 0.00 0.00 0.00 0.00 0.00 0.W 0.03 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.17 0.00 0.00 0.00 0.88 O.oB 0.5, 0.52 0.0-3 0.23 0.34 0.00 0.00 %?B 0.34 0.80 1.03 3.43 2.08 0.23 4.48 1.72 0.40 0.82 0.W 0.W 0.17 0.00 0.W 0.00 0.00 1.03 1.14 0.1, 0.34 0.34 0.02 0.1, 1.37 0.46 O.oB 7.03 ,477 8.42 0.24 0.00 0.00 0.00 0.17 0.00 0.00 0.00 0.68 0.00 0.00 0.04 0.00 0.04 0.W 5.15 0.9, 0.1, 1.20 0.34 0.11 0.48 0.00 0.00 0.04 0.00 0.00 0.00 003 8.18 0.00 0.1, 0.89 0.34 0.00 0.23 0.00 0.00 0.00 0.17 0.0.3 0.00 0.00 0.00 0.00 003 0.00 0.00 0.00 0.00 0.00 0.23 0.00 0.M) 0.M) 0.00 0.00 ‘lo.12 0x3 - 2.30 8.52 1.72 0.34 0.02 1.37
0.00 1.03
gives ID number)
C-l 750 0.57 0.00 0.00 0.18 0.40 0.00 0.00 0.18 0.08 0.W 0.00
0.W 0.00 0.65 0.00 0.00 0.18 0.08 0.00 0.24 0.04 0.4, 0.00 0.08 0.08 0.48 0.08 0.08 0.03 0.W 0.00 0.W 0.04 0.18 0.00 0.00 0.08 0.00 0.00 0.00 0.M) O.MI 0.33 0.33 0.08 0.00 0.16 0.00 0.00 0.00 0.00 0.00 0.W 0.W 0.24 0.24 0.00 0.00 0.04 0.18 0.00 0.04 0.04 0.08 0.00 0.4,
Cl moo
c-1 2wo
c-2 0
c-2 25
c-2 54
0.08 o.Bg o.00 1.83 0.00 0.00 0.00 0.00 0.00 0.w 0.00 0.00 0.15 0.w 0.34 0.00 0.20 5.54 18.88 ,417 0.09 0.00 0.34 2.74 0.00 0.00 0.w 0.00 0.03 0.00 0.00 0.00 0.08 0.00 0.00 0.00 0.00 0.17 0.00 0.M) 0.00 0.W 0.09 0.00 0.00 0.00 O.MI 0.W 0.03 0.00 0.00 0.00 0.00 0.00 0.00 0.00 1.37 0.04 0.M) 0.00 0.00 0.00 0.00 0.00 0.W 0.00 0.17 0.00 0.00 0.34 0.04 0.28 0.M) 0.00 0.00 0.00 0.34 0.03 0.00 0.00 1.83 0.00 0.03 2.08 1.7, 0.48 1.1, O.oB 0.00 0.00 0.00 0.28 0.00 1.38 1.03 0.0, 1.20 0.06 0.00 0.34 0.00 0.5, 0.03 0.M) 0.00 0.04 4.1, 0.60 1.03 5.94 0.00 0.09 0.00 0.00 0.W 2.57 0.12 0.69 2.06 ,.a7 0.00 0.00 0.69 2.40 0.9, 1.28 0.17 0.00 0.00 0.00 0.5, O.og 0.W 0.00 1.37 0.17 0.W 0.00 0.00 1.83 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.09 0.00 0.00 0.28 0.00 0.00 0.00 0.0, 0.77 0.00 0.88 2.40 1.37 0.00 0.W 0.00 0.W 0.34 0.08 0.00 0.00 0.00 0.00 0.03 0.W 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.W 0.00 0.W 0.00 0.04 0.00 0.88 1.03 3.88 0.00 0.M) 0.00 0.W 0.46 0.00 0.04 0.00 0.M) 0.86 0.03 1.38 2.Mi 1.37 0.43 0.W 0.80 2.40 4.1, 0.26 0.00 1.38 0.00 0.46 0.00 0.W 0.00 0.00 0.46 0.28 0.00 0.00 1.37 0.5, 0.08 0.00 0.00 0.00 0.00 0.00 0.00 0.W 0.00 0.00 0.W 2.29 0.W 0.00 0.00 0.03 0.00 0.00 0.00 0.00 0.00 0.00 0.M) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.34 0.00 0.88 O.oB 0.W 0.00 0.00 0.5, 0.17 0.00 0.W 0.00 0.77 0.00 0.00 0.00 0.00 0.00 0.M) 0.00 0.W 0.00 0.52 0.00 0.00 0.W 0.00 0.00 0.00 003 0.00 003 0.00 0.00 003 OS, 0.06 0.00 0.00 2.74 0.00 0.00 0.00 0.00 0.W 0.17 0.08 0.00 1.03 1.83 0.43 0.M) 0.00 0.00 0.48 0.00 0.00 0.M) 0.00 0.04 0.00 0.W 0.W 0.34 0.00 1.80 0.03 1.38 18.80 - 42.00 2.0, 0.00 0.w 0.86 1.54 0.08 0.w 0.5,
Radiolarian
A 0
A 50
A ,@I
12.42 0.00
1.83 0.00 4.58 0.00 0.M) 1,.44 0.00 0.00 0.00 2.28 0.W 0.w 0.04 0.00 0.00 0.00
1.3, 0.33 5.80 0.00 0.33 12.78
2.76 1.38 0.00 1.38 0.00 0.W 0.00 40.01 0.00 0.00 0.00 0.00 0.W 0.00 0.00 15.18 8.28 63.48 0.W 0.w 0.00 0.00 0.W 0.00 5.52 4.14 0.00 0.W 0.00
0.00 3.20 11.44 5.03 0.00 0.82 0.00 0.00 0.04 2.75 0.00 4.12 0.00 0.W 0.00
0.00 0.00 40.01
0.00 0.00 0.49
1.30 2.78 0.W 1.38 2.78
0.00 0.00 0.00 2.29 1.37
0.00 0.00
4.50 0.00
0.04 0.00 0.00 0.W 0.0-3 0.00 0.00 20.04 om 0.00 5.52
0.M) 0.02 0.w 0.00 0.w 2,Ol 0.48 226.90 0.00 2.22 1.83 0.00 0.00 0.48 1.83
8.28 0.09 1.38 0.00 1.38 22.07 1.38 0.00 0.00 2.76 0.00 2.76 0.00 0.00 0.00 - 0.04
0.48 l2.22 0.92 0.W 0.00 0.43 1.37 2.75 0.00 3.20 0.00 0.40 0.00
0.w 0.33 0.04 10.80 0.00 0.65 0.00 0.33 2.05 0.00 0.00 0.20 41.56 2.20 0.00 1.04 0.04 0.00 0.00 0.33 0.33 4.25 0.33 0.22 0.00 0.00 0.00 3.60 0.85 0.00 0.00 2.85 0.00 1.31 0.00 0.04 5.50 0.00 0.00 0.23 10.38 53.67 238.58 0.00 0.32 0.00
A 300 0.18 0.00 0.40 0.00 0.M) 2.12 0.03 0.00 0.W 0.08 0.00 0.09 0.33 0.00 0.00 0.00 0.18 0.33 4.90 1.14 0.00 0.33 0.00 0.00 0.00 0.33 0.10 0.10 0.10 0.49 0.04 0.40 0.00 0.40 0.00 0.18 0.00 0.00 0.00 0.33 0.00 0.00 0.42 3.10 0.00 0.0-3 0.98 1.3, 47.32 0.04 0.85 0.04
c-1
c,
0
25
0.09 0.00 0.w 0.00 0.00 1.38 0.M) 0.M) 0.00 4.83 0.00 0.M) 0.68 0.00 0.00 0.00 0.00 3.45 43.45 10.35 0.00
0.00 0.00 0.89 0.34 0.00 2.40 0.00 0.00 0.00 6.87 0.00 0.00 0.00 0.00 0.W 0.00 0.00 4.01 50.15 15.00
0.00 0.00 0.00 0.00 0.00 3.45 7.59 2.07 0.00 0.W 1.38 0.04 3.45 0.00 0.00 0.00 1.38 0.00 0.00 0.00 0.00 0.00 207 0.00 0.00 0.00 0.00 210.38 0.00 0.09 0.00
0.00 0.00 240 0.00 0.00 0.03 0.34 8.93 1.03 0.00 0.00
c-t 50
0.23 0.00 1.14 0.00 0.00 11.88 0.W 0.23 1.83 2.29 0.00 0.00 0.00 0.00 0.00 0.00 0.00 1.14 7.77 0.01 0.W 0.40 0.23 0.00 0.00 0.23 0.04 8.40 0.W 1.14 0.00 0.00
0.00 0.00 7.56
0.00 1.60
0.00 0.W 0.00 1.03 8.24
0.00 0.03 0.00 1.83 2.74
3.09 0.W 0.00 0.34 3.43 0.00 0.34 0.09 0.00 310.07 0.88
0.40 0.00
1.03 0.00
vertical
Cl lca 0.06 0.w 0.88 0.00 0.00 0.29 0.00 0.04 0.20 0.17 0.00 0.00 0.00 0.00 0.40 0.00 0.00 0.57 1.15 0.17 0.09 0.00 0.00 0.02 0.00 0.00 0.W 1.15 0.00 0.40 0.00 0.00 0.00 0.17 0.00 0.00 0.00 0.1, 0.11
distribution
c-1 200 0.00 0.00 1.54 0.00 0.00 11.15 0.34 0.00 0.17 1.54 0.W 0.17 0.24 0.17 0.34 0.W 0.W 0.89 10.47 1.54 0.00 0.86 0.00 0.M) 3.20 0.00 0.17 0.80 1.03 1.54 0.00 0.51 0.00 1.20 0.W 0.00 0.00 0.34 1.07
0.00 0.22 0.00 0.00 0.00 22.40 30.83 230.80 0.02
0.33 0.00 0.00 0.00
0.40 0.04 0.00 0.00 0.00 0.W
0.00 13.74 0.00 0.00 0.00 0.05 1.26 0.98 0.33 1.04 0.00 l.00 1.04
0.23 0.60 0.40 0.00 0.W 0.00 0.23 2.51 0.00 1.37 0.00 5.2* 0.00
c-1 300 0.w 0.m
2.58 0.17 0.00 0.00 0.W 0.00 0.17 0.17 0.00 0.02 0.00 0.00 0.00 2.58 0.00 0.00 0.24 0.w 0.00 0.86 0.00 0.W 0.W 0.w 0.04 0.W 0.17 0.00 1.55 0.00 0.00 0.00 0.00 0.17
42.83 0.00
1.43 0.00 0.00 0.00 0.00 0.06 0.75 0.W 0.40 0.03 0.34 0.00
2.40 0.34 0.w 0.W 0.M) 0.00 0.24 0.00 0.17 0.00 1.03 0.17
C-l 400 0.w
0.52 0.0.3 0.0.2
0.17 0.00 0.00 0.00 0.00 0.00 0.17 0.00 0.w 0.w 0.17 0.00
229
patterns
0.03 0.00 0.04 0.00 2.10 0.1, 0.11 0.M) 0.34 0.w 0.00 0.00 0.00 0.00 0.00 0.00 0.W 1.48 0.34 0.W 0.04 0.00 0.00 0.40 0.00 0.11 0.W 0.00 0.1, 0.00 0.00 0.00 0.23 0.00 0.00 O.MI
C-l 500 0.00 0.00 0.1, 0.00 0.00 3.86 0.11 0.57 0.M) 0.00 0.00 0.W 0.11 0.11 003 0.04 0.00 0.00 7.2, 0.23 1.72 0.W 0.W 0.22 1.w 0.11 0.w 0.04 0.00 0.1, 0.00 0.23 0.00 1.03 0.00 0.23 0.00 0.23
0.00 0.11 0.03 0.00 0.W 0.00 0.23 0.W 0.w 0.24 0.11 58.81 0.00
1.02 0.00 0.W 0.W 0.40 1.95 0.00 0.1, 1.26 1.14 78.83 0.00
0.57 0.W 0.00 0.00 0.23 0.00 0.00 0.00
0.22 0.11 0.00 0.00 0.23 0.11 0.00 0.34
0.00 0.00 0.00 0.00 0.00 0.00
0.04 0.00 0.00 0.00
0.00 0.23 0.W 0.02 0.03
0.11 0.00 0.34 0.00 0.43 0.11
Cl 750
o.00 0.00 0.00 0.00 0.00 0.24 0.08 0.08 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.W 0.w 0.00 0.04 0.00 0.00 0.00 0.00 0.03 0.00 0.00 0.00 0.02 0.W 0.16 0.04 0.00 0.04 0.08 0.00 0.00 0.08 0.00 0.W 0.04 0.08 0.W 7.00 0.00 0.08 0.03 0.00 0.03 0.00 0.00 0.W 0.W 0.00 0.00 0.00 0.00 0.00 0.60 0.00 0.00 0.00 0.00 0.00
Cl lK!o 0.w 0.w 0.03 0.00 0.04 2.44 0.77 0.80 0.00 0.09 0.M 0.00 0.17 0.34 0.17 0.00 0.00 0.03 2.0, 0.00 0.00 0.00 0.17 0.00
Cl *cm 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.03 0.00 0.00 0.08 0.03 O.MI 0.20 0.00 0.04 0.00 0.12 0.00
c-2
0
25
0.w 0.0-3 0.00 0.W 0.04 O.OS 003 0.00 0.00 0.00 0.M) 0.00 0.00 0.00 0.04 0.04 0.04 0.00 4.85 0.88
0.00 0.00 0.28 0.09 0.51 0.00 0.00 0.00
0.00 0.00 0.00 0.w 0.00 0.W
0.00 0.00 0.00 0.00 0.00 0.00 0.00 4.15 0.09 0.00
0.00 0.12 0.00 0.03 0.00 0.00 0.00
0.00 0.89 0.00 0.80 0.W 0.W 0.W
0.28 0.00 0.00 0.00 0.00 0.00
0.04 0.03 0.00 0.00 0.00 0.00
1.71 0.17 0.00 0.89 1.54 OS.43 0.17 0.77 0.00
0.44 0.00 0.06 0.06 0.00 3.52 0.00 0.00 0.00
0.W 0.00 0.00 0.00 0.00 0.00 2.08
0.00 0.00 0.00 0.00
0.00 0.W 0.00 0.00
0.00 0.43 0.00 0.53 0.00 0.00 0.00 o.*o 0.34 0.09 0.34 0.00
0.00 0.00 0.00 0.00 0.00 0.00 0.w 0.03 0.W 0.00 0.03 0.03
0.00 0.00 0.00 0.04 0.00 0.00 0.w 0.00 0.W 0.043 0.00
0.00
0.02
0.W
0.17 0.00 0.09 0.W 0.20
0.00 0.00
c-2
0.00
0.00 0.w 0.00 0.00 72.00 0.00 0.00 0.00 0.00 0.W 0.00 0.00
1.03 0.00 0.88 0.34 0.0.2 0.W 0.00
C-2 50 0.w 0.w 1.37 0.40 0.00 15.04
0.00 0.00 0.00 0.00 0.00 0.04 ,5.02 4.1,
0.00 0.00 0.W 0.48 0.00 0.05 0.00 0.00 0.00 0.00 0.00 0.40 42.51 2.28
0.W 0.W 0.00 0.00
0.m 0.0, 0.00 0.W
0.00 0.00 0.00 5.42 0.M)
0.w 0.00 O.Ml 8.86 0.w
0.W 0.00 0.00 0.00 3.09 0.00 0.34 0.03
0.W 0.00 0.05 0.00 0.14 0.00 0.00
0.03 0.00 0.W 0.W 0.00
3.42 5.14 O.BO 0.W 0.00 0.00 0.00 0.00 0.00 0.24 0.00 129.80 0.00 0.34 0.00 0.00 0.00 0.00 0.W 0.00 3.08 2.74 0.00 0.00 0.34 0.00 1.37 0.w 0.0-3 0.M) 1.37 0.5,
0.00 5.03 7.77 3.x1 0.00 0.00 0.00 0.00 0.w 1.37 16.00 0.W 149.03 0.00 0.M) 0.00 0.00 0.00 0.00 7.77 5.03 0.40 0.M) 0.00 3.60 0.00 0.00 0.00 13.28 1.03
230
7 3 4 5 6 7 8 8 10 11 13 14 15
16 17 18 20 21 23 24 25 27 28 32 33 35 36 37 38 39 40 41 43 4-I 45 40 47 48 50 51 52
54 55 Ea 57
5% 59 80 82 86 08 Es 70 71 72 73 74 75 78 7, 78 79 90 B1 82 83 a4 E
S. A. KLING
and D. BOLTOVSKOY
c-2 100
C-2 200
CL? 300
c-2 400
c-2 503
G2 750
c-2 1003
c-2 2ooo
F 0
F 25
F 50
0.w 1.63 0.00
1.03 0.03 0.03 0.86 2.40 0.08 0.00 0.00 0.W 0.00 0.00 0.00 0.00 0.1, 0.00 0.00 0.68 0.1,
6.17 0.34 0.00 1.71
0.78 0.00 0.043 0.76 1.53 0.22 0.W 0.11 0.00 0.00 0.W 0.00 0.00 0.00 0.W 0.00 0.11 0.33 0.W 0.22 0.00 1.0s 0.55 1.m 0.03 0.33 0.11 0.22 0.11 0.11 0.W 0.00 0.22 0.11 0.00 0.00 0.00 0.00 0.00 0.1, 0.00 0.11 0.11 0.44 0.11 0.11 0.W 0.00 0.00 0.00 0.00 0.00 0.11 0.1, 0.22 0.44 0.W 0.00 0.11 0.W 0.00 0.33 0.00 0.00 0.00 0.00 0.00 0.55
1.04 0.12 0.12 0.35 1.05 0.00 0.00 0.12 0.00 0.00 0.00 0.00 0.00 0.00 0.04 0.00 0.23 0.23 0.12 0.48 0.00 2.90 0.23 OH 0.12 0.58 0.12 0.00 0.23 0.12 0.00 0.00 0.12 0.M 0.00 0.03 0.03 0.W 0.12 0.23 0.00 0.00 0.98 0.35 0.00 0.00 0.12 0.00 0.M 0.00 0.00 0.00
0.28 0.0, 0.00 0.00 1.m 0.0, 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.14 0.00 0.W 0.00 0.07 0.14 0.0, 0.0, 0.83 0.1, O.sB 0.00 0.83 0.28 0.28 0.00
0.30 0.04 0.00 0.23 0.76 0.04 0.00 0.00
0.07 0.00 0.00 0.2, 0.29 0.03 0.W 0.W 0.02 0.00 0.W 0.00 0.00 0.07 0.00 0.00 0.02 0.00 0.00
3.45 0.00 0.00 0.00 88.29 11.72 0.60 0.00 0.00 0.00 0.00 0.00 0.69 0.00 0.00 14.48 0.00 0.00 I.38 0.W 88.18 1.38 0.00 13.,0 0.00 0.00 122.73 0.00 0.04 0.W 0.59 1.38 2.0, 37.23 0.00 0.00 0.03 0.W 2.07 71.7, 0.00 2.0, 40.68 2.0, 0.00 0.W 0.00 0.69 0.00 0.00 0.00 0.88 0.00 2.0, 0.00 11.03 0.00
2.00 2.00 0.00 0.00 62.48 *.@I 0.00 0.00 0.00 0.00
5.23
3.92 0.00 0.w 0.00 0.W 0.00 0.00 0.00 0.00 0.33 0.33 0.00 0.W 0.08 2.61 0.00 0.33 0.W 2.20 48.04 0.00 0.08 0.00 0.04 1.31 0.08 0.W 0.00 0.33 1.88 0.00 0.03 0.00 0.04 0.00 0.65
0.00 0.W 0.08 1.31 0.33 0.W 0.33 0.00 0.00 0.00 0.33 0.00 0.00 0.33 0.00 ,.$-a 0.03 0.00 0.33 0.00 0.00 0.95 0.00 0.00 0.23 0.00 0.33 10.78
2.40 0.51
0.00 0.1, 1.03 5.13 1.03 0.5, 0.00 0.34 0.5, 0.24 0.00 0.00 0.1, 0.08 0.00 0.17
0.00 0.00 0.17 0.00 0.00 0.00 0.17 0.51 0.51 0.04 0.W 0.00 0.00 0.34 0.04 0.00 0.1, 0.00 a.@3 2.22 0.00 0.00 1.7, 0.M 0.00 0.1, 0.17 0.17 0.00 0.00 0.M 1.e3
5.48
0.00 1.03 0.1, 0.00 0.00 0.1, 0.00 0.1, 1.3, 0.51 0.51 0.5, 0.04 0.51
0.60 5.88 0.51
2.23 0.24 0.00 0.34 0.51
0.00 0.00 0.09 1.03 0.W 0.51 0.00 0.00 0.04 0.00 0.00 0.00 0.88 0.34 0.03 0.1, 0.17 0.W 0.00 0.00 0.17 0.1, 0.@8 1.20 0.00 0.00 2.40 0.W 0.00 0.51
0.17 0.34 0.M 0.W 0.17 3.28
0.00 0.23
0.04 0.58 030 0.00 2.20 0.00 0.00 0.12 0.00 0.11 0.00 0.00 0.00 0.93
0.0, 0.0, 0.00
0.W 0.21 0.00 0.0, 0.0, 0.0, 0.w 0.00 0.00 0.0, 0.42 0.21 0.00 0.14 0.00 0.04 0.w 0.W 0.00 0.00 0.04 0.14 0.00 0.21
0.00 0.00 I.40 0.00 0.09 0.14 0.0, 0.00 0.07 0.00 0.14 0.0,
0.04 0.00 0.00 0.04 0.0-l 0.04 0.00 0.00 0.w 003 0.08 0.04 0.04 0.11 0.08 0.00 0.23 0.W 0.04 0.08 0.00 0.00 0.00 0.00 0.04 0.04 0.00 0.00 0.00 0.00 0.04 0.W 0.00 0.19 0.04 0.04 0.03 0.00 0.08 0.00 0.00 0.00 0.00 0.04 0.08 0.00 0.04 0.M 0.00 0.48 0.00 0.00 0.11 0.00 0.04 0.04 0.04 0.0-2 0.23
0.15
0.00 0.12 0.02 0.38 0.05 0.34 0.00 0.0, 0.w 0.05 0.00 0.w 0.05 0.07 0.00 0.02 0.00 0.W 0.00 0.00 0.00 0.W 0.07 0.02 0.05 0.00 0.00 0.05 0.00 0.00 0.00 0.00 0.02 0.05 0.02 0.28 0.00 0.00 0.0, 0.M 0.00 0.05 0.00 0.W 0.00 0.00 0.00 0.05
0.00 0.00 0.00 0.w 0.00 2.08 0.03 0.00 0.0s 0.00 40.51
0.04 0.00 6.8, 0.w 0.69 70.03 0.00 0.00 0.03 0.00 0.00 0.00 35.01 2.75 0.02 0.00 0.00 0.00 58.04 0.00 0.00 24.72 1.3, 0.69 0.00 0.00 0.w 0.00 0.00 0.00 0.00 0.00 8.57 0.09 2.75 0.00 4.12 0.04 0.W 2.06 3.43 0.00 0.00 2.78 1.37 3.45 2.08 0.00 0.w 2.78 2.08 30.34 8.18
F 100
F .mo
1.37 0.00 0.00 0.00 35.18
4.14 0.00 0.00 3.11 23.82
0.86 0.w 0.w 0.52 13.57
8.23 0.00 0.00 0.00 0.00 0.w 0.00 0.w 0.00 0.00
1.04 0.52 0.00 0.w 0.w 0.00 0.00 1.04 0.00 0.52 0.52
0.00 0.00 0.00 0.04 0.00 0.00 0.00
0.17 0.00 0.w 0.00
0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 2.58 0.W 0.00 1.03 0.00 0.00 0.52 0.52 0.W 0.34 0.1, 0.00 0.00 2.58 0.00 0.00 0.W 0.00 0.00 1.03 0.04 0.00 1.72 0.17 0.88 0.00 0.00 0.03 0.00 0.34 0.00 0.04 0.00 0.00 0.00 0.34 0.1, 0.00 0.00 0.00 0.04 0.17 0.00 0.5S 0.1, 0.00 0.W 1.03
0.00 0.00 0.W OS-XI 0.00 0.17 0.00 0.00 3.00 0.5, 0.86 1.88 1.03 0.00 0.60 0.1, 0.00 0.00 0.00 0.00 0.00 0.88 0.00 0.00 0.00 0.00 0.00 2.5, 0.00 0.00 0.68 0.1, 0.34 0.00 0.1, 0.34 0.W
2.74 0.00 0.00 0.03 0.00 21.02
0.48 0.00 2.23 0.00 0.01 39.30 0.00 0.00 0.40 0.00 0.00 0.w 15.08 0.00 0.04 0.03 0.03 0.00 39.30 0.W 0.91 15.54 4.5, 0.46 0.00 0.04 0.00 0.00 0.04 0.40 0.02 0.00 133 0.00 1.3, 0.00 0.00 0.00 0.47 0.91 1.83 0.00 2.29 1.M 0.00 0.00 1.37
0.00 0.00 0.52 0.00 24.88
0.w 0.W 25.80 0.04 0.W 6.73 0.04 0.52 0.52 0.52 0.00 0.52 5.70 0.04 0.00 0.00 0.00 0.W 8.80 0.52
0.03 3.11 1.04 8.80 0.00 0.52
0.00 0.00 0.00 0.00 0.00 0.02 0.00 0.00 2.58 0.00 1.04 0.52 0.W 1.04 2.0, 0.00 0.00 0.00 0.00 0.00 13.88
F 3w 1.37 0.w 0.00 0.00 0.77
0.34 0.00 0.00
0.51
0.00 0.04 0.00 0.04 0.W 1.37 0.00 0.17 1.37 0.1, 0.1, 0.1, 0.04 0.51 0.17 0.W 0.04 1.03
F 400
F 500
F 750
F loco
0.23 0.48 0.00 0.23 5.76 0.12 0.w 0.00 0.00 0.00
0.34 0.11
0.00 0.03 0.00 0.03 l.w 0.00 0.00 0.00 0.04 0.00 0.00 0.00 0.00 0.00 0.00
0.w 0.0s 0.00 0.23 1.57 0.00 0.03 0.00 0.w 0.0-3 0.w 0.00 0.00 0.20 0.00
0.00 0.03 0.00 0.00 0.00 0.03
0.06 0.06 0.00 0.00 0.00 0.11
0.W 0.00 0.00 0.W 0.00 0.04 0.00 0.23 0.00 0.00 0.81 0.48 0.35 1.50 0.w 0.23 0.58 0.00 0.00 0.12 0.03 0.00 0.00 0.35 0.00 0.00 0.00 0.00 0.00 0.02 0.00 0.00 0.00 0.35 0.35 0.12 0.00 0.12 0.00 0.00 0.00 0.W 0.00 0.23 0.12 0.58 0.12 0.12 2.89 0.00 0.12 0.00 0.00 0.04 0.00 0.04 0.00 0.23
0.48 3.30 0.00 0.00 0.00 0.00 0.00 0.00 0.11 0.00 0.17 0.00 0.00 0.00 0.W 0.57 1.48 0.11 0.34 0.00 0.00 0.23 0.17 0.11 0.1, 0.00 0.04 0.w 0.11 0.00 0.00 0.00 0.00 0.5, 0.W 0.00 0.40 0.23 0.11 0.00 0.08 0.00 0.00 0X-J 0.W 0.00 0.W 0.00 1.08 0.W 0.21) 0.00 0.00 0.00 0.00 0.06 0.08
0.23 _s 0.26 0.08 0.14 0.20 0.86 0.00 0.14 0.00 0.W 0.00 0.23 0.00 0.1, 0.08 0.W 0.00 0.00 0.W 0.W 0.00 002 0.00 0.03 0.00 0.17 0.00 0.04 0.00 0.00 0.08 0.09 0.00 0.00 0.03 0.00 0.11 0.20 0.00 0.00 0.00 0.00 0.1, 0.34 0.00 0.03 0.00 0.03 0.00 0.00 0.W 0.00 0.00 0.03 0.00 0.W 0.03 0.W 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.14 0.03 0.00 0.23 0.00 0.00 0.00 0.00 0.1, I.04 0.00 0.03 0.W 0.03 0.03 0.1, 0.00 0.00 0.08 0.03 0.00 0.M 0.00 0.08 0.00 0.03 0.09 0.09
Radiolarian
vertical
distribution
231
patterns
c-2
c-2
c-2
c-2
c-2
c-2
c-2
c-2
F
F
F
100
m
300
4m
Em
750
loo0
2Mx)
0
25
53
0.00 0.51 0.w 0.00
0.w 1.03 0.00 0.00
F too
F 200
F 3w
F 400
F 5ca
F 750
F 1oocl
0.w 0.00
0.w
0.03 0.00 0.w
8.57 0.34
0.87 0.11 0.w 0.00 0.00 0.00 0.44
a00 2.07
0.34 0.00
0.00 0.08
0.00 cLcK7
0.w CKO
0.w 0.00
0.w 0.00 0.51 0.17 0.w 0.17
0.17 0.00
0.00 0.00 0.17 0.00
0.22 0.00 0.00 0.00
0.w 0.06
0.w cl.03 0.00 0.44
0.w 0.23
0.w 0.00 mm 0.14
Deep-Sea Research I, Vol. 42, No. 2, pp. 191-231. 1995 Copyright 0 1995 Elsevier Science Ltd Printed in Great Britain. All rights reserved 0967437195 $9.50 + 0.00
0967-0637(94)00038-7
Radiolarian vertical distribution patterns across the southern California Current STANLEY A. KLING* (Received
29 August
and DEMETRIO BOLTOVSKOY~
1993; in revisedform
5 July 1994; accepted
8 July
1994)
Abstract-Polycystine radiolarians were identified in 36 plankton samples collected at depths ranging from 0 to 2000 m at four stations extending west from about the U.S.-Mexico border (approx. 32”N, 117”W to 124”W), in November-December 1977. In total, 136radiolarian taxa were recorded, but 90% of all individuals were accounted for by only 40 of these. Highest abundances were found either at the surface, or at 25-50 m. Based on maxima in the vertical profiles of the most abundant radiolarians, three major depth-intervals were defined in the upper 300 m: O-50 m, 100 m and 200-300 m. Between-station-similarities in the specific makeups of these layers, however, were low. Thirty-nine taxa had peak abundances below 300 m at one or more stations; 11 of these are probably deep-water forms. Although in terms of individuals per liter of water filtered, upper-layer taxa are noticeably more abundant than deep species, the latter have much more extended depthranges, which might significantly enhance their sedimentary output. The inshore and oceanic stations shared very similar, warmer-water radiolarian assemblages in the uppermost 25 m, whereas the intermediate station was dominated by colder-water forms at those depths. Below 50 m, however, the inshore station had enhanced proportions of deeper- and colder-water species, differing strongly from the oceanic site. We suggest that this pattern results from circulation of the Southern California Eddy, which transports Central Water from the oceanic station on the western edge of the California Current around the intermediate stations to the inshore stations. The coldwater signal at subsurface layers of the inshore station could be reinforced by coastal upwelling and southward transport by the California Current, thus further enhancing the abundances of deeperwater radioiarians at this site. Analyses of the effect of such vertical patterns on paleoceanographic interpretations stress the importance of the signal of “environmentally neutral” deep-living species, as well as that of shells produced in the near-surface layers of distant areas and transported at depth to the region of the study.
INTRODUCTION THE overwhelming
majority of information on polycystine biogeography has been derived from surface sediment samples, whereas data on the distribution of live assemblages, and especially on species-specific vertical distribution patterns, are very scarce and largely restricted to a few oceanic areas (e.g. RENZ, 1976; MCMILLEN and CASEY, 1978; KLING, 1979; MORLEY and STEPIEN, 1984; DWORETZKY and MORLEY, 1987). Such information, however, has an important bearing on current problems associated with the fate of organic matter generated in the upper mixed layer (e.g. GOWING and GARRISON, 1992). Analyses of vertical distribution patterns can also yield valuable data on the modes of sedimentation
“416 Shore View Lane, Leucadia, CA 92024, U.S.A. ?Departamento de Ciencias Biologicas, Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires, 1428 Buenos Aires, and CONICET, Argentina. 191
192
S. A. KLINC
and D. BOLTOVSKOY
of radiolarian shells and help assess the relative importance of dissolution en route to the sea-floor and of destruction by grazing (MILLIMAN and TAKAHASHI, in press; BOLTOVSKOY and ALDER, 1992a). Many paleoceanographic studies based on comparisons between surface sediment distributions and downcore sequences employ analytical techniques (e.g. factor analysis and transfer function techniques: IMBRIE and KIPP, 1971) that assume irrelevance of living organisms in the overlying plankton. However, these assumptions do not take into account biases associated with vertical distribution patterns of such deepliving groups as radiolarians. In addition, at any given locale bottom thanatocoenoses are made up of locally produced shells and, to different extents, by shells produced elsewhere and transported laterally via subsurface and deep currents (BURCKLE, 1981; POKRAS and MOLFINO, 1986; BOLTOVSKOY, 1988, 1991, 1992; BOLTOVSKOY and ALDER, 1992a). Thus, climatic oscillations which affect the strength of these currents (and, consequently, the intensity of advection of sedimenting shells) can introduce bias into direct comparisons between recent and past geographic distribution patterns. The present work describes the vertical (O-2000 m) distribution of polycystine Radiolaria at three closely spaced sites off the Southern California coast dominated by a strong gradient in physical and biological variables. In addition, duplicate sampling at one of the sites provides information on the temporal (approx. one week) variability in polycystine makeup and distribution. Although the patterns analysed are but a snapshot image of a highly variable situation, they nevertheless offer clues for the interpretation of the ecological settings of the radiolarians involved and, by extension, on their paleoceanographic significance in the fossil record. MATERIALS
AND
METHODS
Samples were collected from a line of stations extending west from about the U.S.Mexico border (Fig. l), as part of BMET Cruise in late November and early December of 1977. Station data are as follows: Sta. A: 23 Nov. 1977,32”38’N, 117”29’W; Sta. Cl: 25-27 Nov. 1977,32”30’N, 120”26’W; Sta. C2: 30 Nov.-3 Dec. 1977,32”38.25’N, 120”35.75’W; and Sta. F: 5-6 Dec. 1977,32”38.23’N, 123”53.96’W. Field and laboratory methods for this study are essentially the same as those described by KLING (1979). Plankton samples were obtained with opening-and-closing nets controlled and monitored by an acoustically operated system described by BAKER et al. (1973). The nets, made from Nitex nylon mesh with 62-pm opening, were designed to present a 1 m by 1 m cross section to the towing path while being towed horizontally with the opening frame at a nominal 45 degree angle. They were about 3.5 m long midway along the tilted vertical frame members. Flow-meters were not available during the tows reported here, so volumes of water filtered were estimated on the basis of distance towed, calculated from ship speed and time, and net mouth diameter (1 m2). Earlier experience with the same net system equipped with an external flow meter indicates that distance calculated from ship speed and time is a reliable measure of distance travelled. Correlation between metered and calculated distances were very highly significant (Y = 0.999, P < O.OOl), which suggests that our distance estimates are generally correct. Nevertheless, this ignores the effects of net clogging, which are admittedly serious despite the relatively large size of the nets (1 m2 x 3.5 m) and the mesh openings (62,um). To compensate in some way for this shortcoming, the length of tows was much shorter in
Radiolarian vertical distribution patterns
193
the upper, plankton-rich layers (on the order of 100 m at 0 m, 200 m at 25 m) than those in deep waters, where particulate materials are scarce (700 m at 500 m, 1500 m at 1000 m, 3000 m at 2000 m). This does not ensure that the values given are correct, but it does lend support to the contention that they are at least internally consistent. We have attempted to minimize the effects of water volume inaccuracies upon the reported analyses in two ways. First, where we used numbers of individuals per cubic meter, for example to identify abundance maxima (Table 2), we considered only values that exceed background levels by about five times or more (see the vertical profiles summarized in Fig. 7). Second, most of the statistical calculations were based on data transformed into percentages. In the laboratory, samples (preserved in formaldehyde) were split and one fraction combusted to rid it of entangling soft tissue. Measured subsamples of the combustion residues were transformed to cover glasses and dried. Cover glasses were mounted on glass slides with Pleurax. All specimens on a cover glass were identified and counted. Unidentified forms were lumped in a separate category. Counts in all categories were converted to numbers per m3 of water filtered. Unidentified radiolarians, including juveniles and incomplete shells, made up very large proportions of the overall assemblages retrieved: 30.7-66.7% (see Table 1). These individuals were only included in analyses of total radiolarian standing stocks. Speciesspecific distributional interpretations, especially those based on percentage data, are obviously affected by these high proportions of unidentifieds, yet the bias seems to be evenly distributed over the entire database, since no relationship was found between the percentages of unidentifieds and site or depth. The number of individuals per sample scanned varied between 156 and 2409 (Table 1). Despite this wide range in sample-size, the database seems to represent adequately the overall specific diversity of the area insofar as no correlation was found between the numbers of specimens identified and the numbers of species recorded (1. = 0.173, P > 0.1). OCEANOGRAPHIC
SETTING
The transect of stations for this study extends across the California Current (REID et al., 1958) from the Southern California Borderland to the deep Pacific basin (Fig. 1). Station A was over the San Diego Trough with a bottom depth of about 1000 m. Stations Cl and C2 were located just west of the Patton Escarpment where water depths exceed 4000 m. Station F was in the deep ocean where water depths are in the 5000-m range. Hydrographic casts were made at each station and the resulting O-300 m temperature and salinity profiles are presented in Fig. 2. Because of the equipment failures, no salinity data are available for Stas C2 and F. The temperature profiles indicate that the surface mixed layer extends to a depth of about 100 m at the western Sta. F, shallows to about 50 m at the central Sta. C2 and 25 m at Sta. Cl, and shallows further to or near the surface at the inshore Sta. A. Salinity profiles are marked by a shallow salinity minimum at about 50 m at Sta. Cl and A (where data are available). A shallow salinity minimum is a regional feature of the California Current (REID et al., 1958;REID, 1973). The deeper salinity minimum of the North Pacific Intermediate Water (REID, 1965) is typically not maintained in the California Current, and does not appear in these profiles. The major component of flow in the California Current is southward, However, a subsurface countercurrent is also a prominent feature, and at times the countercurrent
194
S. A.
KLING
and D.
BULTOVSKOY
Table 1. General data for the materials surveyed. Equitability (E = HE&,,,, = @(Pi.logz Pi) + Ps.logz Ps); where Pi is the proportion of the ith species, and s is the total number of species in the sample; PIELOU, 1966), is inversely proportional to the degree of numerical dominance of one or a few species over the rest of the assemblage, whereas specific diversity (H’ = Z Pi .log2 Pi, where Pi is the proportion of the ith species; SHANNON and WEAVER, 1949)) increases with increasing equitability and species richness
station
Total individuals
Depth
bd.
m-3
counted
I ,4
300-1000
Specific diversity
Equitability
No. of spe
1661 966 521 608 111 121 64 37 9 21
42.45 40.27 40.56 52.98 56.97 49.86 54.14 53.23 51.47 46.75
4.734 4.544 4.544 4.771 4.289 5.023 4.881 4.983 4.460 5.146
0.786 0.801 0.790 0.792 0.790 0.854 0.837 0.834 0.863 0.842
65 51 54 65 43 59 57 63 36 69
665
47.18
4.651
0.8M
56
588
50
51.09
4.899
0.846
57
50 UN 200
156 700 910 1227 596
108 240 416 401 102
66.67 54.00 35.82 60.15 44.20
4.224 4.171 4.383 3.843 4.632
0.899 0.801 0.789 0.692 0.791
26 37 47 47 58
300 400 500 750 1000 2000
706 284 328 286 291 316
121 31 38 20 11 8
42.41 44.01 41.46 40.21 51.55 49.05
5.042 5.155 5.007 5.013 5.068 4.777
0.831 0.909 0.870 0.871 0.893 0.875
67 51 54 54 51 - 44
716
231
so.54
4.382
0.800
47
379
44
43.93
5.057
0.875
55
304
9
50.30
4.923
0.884
48
622 1843 1925 1673 2347
429 633 440 96 403
49.04 50.03 52.47 64.67 48.53
4.319 4.494 4.772 4.602 5.241
0.812 0.746 0.795 0.782 0.816
40 65 64 59 86
594 782 1311 319 1308 275
102 89 150 26 112 8
42.09 63.94 52.56 30.72 50.42 44.00
4.732 4.353 5.037 3.466 5.209 4.834
0.804 0.771 0.814 0.643 0.842 0.880
59 50 73 42 73 45
1501
351
51.14
4.693
0.793
62
863
96
47.94
4.559
0.775
59
792
60
47.21
5.022
0.861
59
822
1134
44.18
4.559
0.812
49
1003 1751 766
459 573 125
51.77 42.21 37.91
5.031 4.745 5.067
0.845 0.762 0.816
62 75 74
1086
573
44.02
4.850
0.809
65
m mean
O-300 m mean 300-1000 m mean 1000-2000 m mean
300-1000
bldh.
2409 1407 1140 1772 646 706 556 650 307 723
0 2s
c2
% unid.
m mean
Radiolarian
vertical
125
distribution
195
patterns
120'
115 35"
32"
Fig.
1.
Fig. 2.
Sampling
Vertical
locations
and November (O-500 db),
Temperature
(“C)
10
15
O-300
195C1964 after WYLL~E
mean geostrophic (1966).
flow
Salinity
to\-)
m profiles of temperature and salinity at the stations shows profiles down to the deepest layers covered.
surveyed.
at the surface
Inset
graph
reaches the surface. The surface countercurrent is particularly well developed close the coast in southern California and in winter months. These opposing currents define a semipermanent circulation system (the Southern California Eddy) over the Borderland in the broad embayment of the southern California coastline south of Point Conception to Punta Baja in Baja California (the Southern California Bight). As a result, the central part of the region may differ hydrographically (e.g. lower temperatures and salinities) from the waters either farther onshore or offshore. Similarly, inshore waters may be more closely
196
S. A. KLING
and D. BOLTOVSKOY
related to the far offshore Central Water Mass than to the core of the California Current. We note in our data that surface temperatures are slightly lower at the central stations (Cl, C2) than at the outer stations (A, F) (Fig. 2). Available surface salinity data are consistent with the temperature observations: lower at Sta. Cl than at A. The California Current is a very complex system that changes rapidly in space and time. However, PELAEZ and MCGOWAN (19S6) recognized a number of major features in the southern California Current that.appear to have both hydrographic and biologic significance. The structures were identified from satellite images of phytoplankton pigment concentrations near the ocean surface. Three features are of interest to this study. An area of high pigment concentration is probably related to the Southern California Eddy and located over the bathymetric high area of the outer Borderland basin system (the location of Stas Cl and C2). Secondly, a low-pigment intrusion flows from the south in the countercurrent close to the coast off San Diego in the region over the deep San Diego Trough (Sta. A; see Fig. 2), and is especially well-developed in summer and fall months (July-December). This low-pigment water is related to a third feature, the low-pigment, oligotrophic waters of theNorth Pacific Cental Gyre to the west (Sta. F), which circulate around the southern and eastern borders of the high-pigment area. Although in detail these patterns shift position, expand and contract, and change in pigment concentrations with time, they were always identifiable in some form. A seasonal cycle in their development noted by PELAEZ and MCGOWAN (1986) were repeated in the three years’ records they analysed. RESULTS AND DISCUSSION Specific makeup
In total, 136 radiolarian taxa were identified in the collection (see Appendices 1,2). This number compares favourably with previous studies of complete polycystine assemblages in the California Borderland. BOLTOVSKOY and RIEDEL (1987) recorded 155 species in forty-eight O-100 m plankton tows along the California Current; and CASEY (1966) reported approx. 200 species from the same area between October 1962 and February 1964. However, as in previous studies, a rather small proportion of these 136 taxa was represented by common to abundant forms, with the bulk being recorded sporadically and in very low numbers. Thus, when the entire database is considered, only about 40 radiolarians accounted for ca 90% of all identified individuals. Although the rare species do convey important ecological information, probably even more than the abundant ones (e.g. BOLTOVSKOY, 1989), their scarcity poses serious methodological problems to interpretation of associated environmental signals. In fact, absences of these rare organisms from the samples are questionable, rather than presences, because of the impossibility of distinguishing the combined effects of ecological exclusion and insufficient sample-size. The only practical way to overcome this problem is to restrict the database to those species whose absences can be reasonably assumed to result from environment-related causes (rather than to procedural artifacts). Therefore, we based many of our subsequent detailed analyses on a reduced set of 41 taxa with at least 20 individuals in one or more samples. This arbitrary cutoff value seems an acceptable compromise which retains a moderately large inventory, yet excludes the questionable absences. These 41 abundant radiolarians comprise 6491% of the entire inventory of each sample (excluding the
Radiolarian
vertical
distribution
197
patterns
‘50 100 300
n
cl
Fig.
3.
Vertical
distribution
of total radiolarian
concentrations.
unidentified specimens). However, in agreement with the pattern of total abundances (Fig. 3), their relative contributions were higher in the top layers than at depth.
Interpretation
of the vertical profiles: in situ living populations vs dead sedimenting shells
A major problem for the assessment of radiolarian depth ranges is the lack of information on the physiological status of the individuals (i.e. live cells or dead, sinking skeletons). It is well known that, although specimens retrieved in the uppermost layers represent mostly live organisms, the proportions of empty shells increase significantly with increasing depth (e.g. PETRUSHEVSKAYA, 1971; TAKAHASHI, 1983/1984; GOWING and COALE, 1989; BOLTOVSKOY et al., 1993a). In our materials concentrations below the uppermost maximum decrease either more or less gradually, or abruptly, but most profiles suggest a restricted preferred depth range, with a “tail” made up of “stray” live (?) specimens and settling dead individuals (Fig. 3). In some instances, however, the profiles show several maximum throughout the water-column (e.g. Sta. Cl, see Fig. 3). These patterns are amenable to alternative interpretations. Although the upper peak is probably due to live individuals, the lower ones can be: (1) enhancements due to favourable local conditions for some of the taxa concerned; (2) accumulations of skeletons transported laterally from elsewhere; (3) sinking accumulations resulting from short-lived blooms at shallower depths; (4) concentrations of sinking shells derived from discontinuities in the density profile of the water-column, or a combination of the above. The secondary peaks below primary ones higher in the water-column largely represent settling shells, rather than in situ blooms, is suggested by a close analysis of the four maxima in the profile of Sta. Cl (at 25, 200, 500 and 1000 m, cf. Fig. 3). Although the uppermost peak (at 25 m) is totally dominated (91%) by taxa that are at maximum abundance precisely in this particular stratum, the dominance of locally peaking species
198
S. A. KLINC andD. BOLTOVSKOY Ten
25 m species
Two
200
m species
Three
500
m species
Three
1000
m species 3
T. octacantha E. acuminarum P. t&bum D. tetrathalamus 5. irregularis L. quadran u/a Perrdium (?P sp. P. praetextum T. trachelium P. hertwigii
tetras
1800 1 zooo-
davisiana
L. buetschli cornutoide
II
0
80
160
10
20
30
Individuals
L
Spongorus sp. T. bicornis f?) sp. 1
Carpocanium
Dicryophimus
per
0
1
sp. 2 2
m-3
Fig. 4. Vertical profiles of the species characteristic of the four peaks in total radiolarian abundance at Sta. Cl (cf. Fig. 3). In proportion of total individuals, these taxa were 22 times more abundant at the given layer than at any of the other three layers compared.
decreases to 63% at 200 m, 20% at 500 m and 25% at 1000 m. Conversely, the contribution of species presumably generated elsewhere (i.e. peaking above a given depth), is 32% at 200 m, 66% at 500 m and 75% at 1000 m. The remainders, however, are made up of taxa that are at maximum abundance within the given layer, and therefore probably represent live populations, or at least assemblages not generated in the upper strata of the same locale (see below). Figure 4, based on a selection of the most reliable data points, shows that the uppermost (25 m) maximum at Sta. Cl is dominated by 10 clearly surface-dwelling radiolarians (52% of the overall assemblage at this depth). These taxa account for 14,6 and 10% at 200,500 and 1000 m, respectively. Theocalyptra davisiana cornutoides and Larcopyle buetschlii account for 19% of all identified radiolarians at 200 m, although they represent only 1% at 25 m, and 4-5% at 5OGlOOO m. Spongurus sp., Theocalyptra bicornis and Dictyophimus sp. 1 comprise 17% of the (identified) individuals at 500 m, whereas they are absent altogether at 25 m, and represent 0.7 and 4.5% of the radiolarians at 200 and 1000 m, respectively. Finally, Siphocampe arachnea, Porodiscus rnicroporus and Carpocanium sp. 2 are noticeably more abundant at 1000 m than elsewhere. Although their overall contribution to this layer is a low 4%) they represent only O-0.6% of the assemblages at 25, 200 and 500 m. Figure 4 also shows that minor yet noticeable peaks for the taxa characteristic of the strata reviewed also occur at some of the levels of maximum total radiolarian abundance (cf. Fig. 3). Th us, the ten 25 m species also have high concentrations at 200 m; the two 200 m ones at 25 and 500 m, etc. Although secondary maxima above the layer of maximum abundance are most probably due to live individuals (probably the initial part of the downward abundance increase), it seems difficult to envision that a species that thrives, for example, in the upper 50 m can again find very favorable conditions at 200-1000 m. The percentages of Fig. 4 are based on total radiolarian absolute abundances, for which our
Radiolarian
vertical
distribution
199
patterns
Total species recorded
Species added (Oh of total)
Fig.
5.
Total number new species
of radiolarian species added to the inventory
recorded in each sample (top panels), and numbers of the overlying samples (bottom panels).
of
estimates are somewhat rough (see Materials and Methods). Thus, biases in these numbers, on which concentrations of individual species are based, greatly affect the shapes of the profiles shown. That radiolarian assemblages at depth increasingly represent flux of skeletons out of the upper layers is also strongly supported by the fact that neither the number of taxa, nor specific diversity or equitability, vary systematically with depth (Table 1). Figure 5 illustrates the numbers of species recorded in each sample (top panel), and the new species added to the total inventory of each station as one progresses from top to bottom (bottom panel). It is noteworthy that, although most of the taxa are present at all depths, deeper samples contribute successively smaller numbers of new taxa to the roster encountered in the overlying waters. The upper 50 m account for 55-72% of the total diversity of each site (Fig. 5). Interestingly, the higher vale (72%) is that of Sta. Cl, which showed the most pronounced subsurface abundance peaks (Fig. 3). This reinforces the above-discussed premise that these deep maxima at 200,500 and 1000 m are either biases in our estimates of total radiolarian abundance, or accumulations of dead skeletons.
200
S.A.KLING~II~D.BOLTOVSKOY %
of
“live”
Polycystines
“Live” polycystines (regression vs. log depth, r = - 0.897) % of “live” polycystines (actual data points)
-llIlTota’ 5,
polycystines per m-’
polycystines
per
0
Total
Fig. 6. Average percentage of “live” polycystines and mean total radiolarian shells vs sample depth;
m-3
(see text for description of assumptions used) based on combined data for entire collection.
We also attempted to address the problem of the differentiation between living and dead radiolarians by assuming that individuals retrieved at and above the level of a species’ maximum abundance were live cells, and that all those below the maximum were dead skeletons settling through the water column. For all taxa “live” and “dead” specimens were summed separately, and the average ratio between the two figures was calculated using all four sites samples. Figure 6 illustrates the curve thus obtained, superimposed on the regression of percentage of “live” individuals on log depth (meters + 1). The assumptions used in this approach have some obvious pitfalls: (1) a species can peak at more than one depth-horizon; (2) downward extensions of the preferred living depth-range are ignored and attributed to settling shells; and (3) local accumulations of sinking skeletons can be mistakenly taken as preferred living depths. Nevertheless, interestingly, these results agree remarkably well with a similar analysis of the vertical distribution of live and dead radiolarians based on the presence of protoplasm in their shells. In the equatorial Atlantic about 2% of the polycystines collected in sediment traps at 850 m are individuals with protoplasm (BOLTOVSKOY et aE., 1993a). It should be emphasized that differentiation between skeletons with and without protoplasm in vertical plankton profiles can furnish clues on the living depth ranges of the species, yet it does not provide unequivocal information because of the uncertainties associated with the speed of decomposition of the protists’ cytoplasm. BOLTOVSKOY and LENA (1970)) for example, concluded that specimens of several planktonic Foraminifera
Radiolarian vertical distribution patterns
201
still contained protoplasm in their shell 98 days after death. This lapse is significantly longer than the time it takes a radiolarian shell to reach the sea-floor (TAKAHASHI and HONJO, 1983). Information on living depth ranges vs export ranges are also intimately linked with processes responsible for the production of new shells, and with the sinking speeds of the skeletons, For surface-dwelling taxa, for example, comparison of standing stocks in the upper layers with concentrations at depth can furnish hints on reproduction rates. In our database, the mean abundance of the 20 radiolarians which peaked at 0 m at Sta. F (see below) was approx. 346 ind m -3 in the O-100 m stratum (which, presumably, is where the majority of these species reside). Between approx. 600 and 1000 m their concentration was rather stable, averaging 3.4 ind. rnm3. Thus, assuming a sinking speed of 1.50 m day-’ (TAKAHASHI and HONJO, 1983), losses by sinking from O-100 m, as recorded by our 60s 1000 m samples, are 1.47% of the population at O-100 m. This translates into a turnover rate of 68 days. This estimate is somewhat higher than previous (indirect) estimates of radiolarian reproduction rates, which suggest values ranging around 20-30 days (see review in CARON and SWANBERG, 1990). Faster sinking speeds and shallower living ranges for the taxa considered, among other factors, would increase this doubling rate. A major point, however, is most probably the loss of shells due to destructive grazing by larger zooplankters. BOLTOVSKOY and ALDER (1992a), for example, concluded that over 90% of the radiolarian shells produced at O-400 m in the Weddell Sea are destroyed beyond recognition above 900 m. Our results suggest that, if radiolarian life-spans are effectively around 20-30 days, in the area of the study destructive grazing eliminates 30-50% of the output from the upper 100 m. It should be stressed that the impact of this process is likely quite uneven on radiolarian taxocoenoses, more fragile skeletons being eliminated in larger percentages than the more robust ones. Comparison of sediment trap vs surface sediment assemblages in the eastern equatorial Atlantic (BOLTOVSKOY et al., 1993b) suggest that the relative contribution of spongodiscid shells might be over-represented in the “grazed assemblages”, whereas the nassellarians, and especially their young, developing forms, might be under-represented. Based on the above considerations, our analyses of the vertical distribution patterns and depth preferences of individual radiolarian species paid special attention to the uppermost layers of peak abundance of each taxon. Vertical distribution patterns Highest radiolarian abundances were recorded either at the surface (Stas A, F), or at 25 (Sta. Cl) to 50 m (Sta. C2). Secondary maxima below this upper layer were generally minor, with the exception of Sta. Cl, where samples from 200 m, and, to a lesser extent, from 500 to 1000 m, presented noticeable peaks in radiolarian concentration (Fig. 3 and Table 1). Because most of the radiolarians were concentrated in the upper most layers, and in order to facilitate comparisons between Sta. A (which was sampled to 300 m only) and the rest of the sites surveyed, detailed treatment of vertical patterns was subdivided into two depth-intervals: O-300 m, and O-2000 m (for the species which showed major peaks below 300 m only). In the O-300 m section a total of 38 species was included (of the 41 for which over 20 specimens were recorded in at least one sample, Spongopyle osculosa, Larcopyle
202
S. A.
KLING
and D.
BOLTOVSKOY
buetschlii and Spongurus sp. were eliminated because they had deep distribution patterns). In the O-2000 m description we considered all the taxa that peaked below 300 m in one or more sites. The upper 300 m
In general terms, species-specific vertical profiles varied widely between stations. As a first approach to the assessment of congruence between sites, we correlated all station pairs of vertical O-300 m profiles of the 41 most abundant radiolarians. Only 16% of the indices thus obtained were significant (positive) at the 95% level, and about twice as many (33%) were negative. Surprisingly, the lowest proportion of significant (P < 0.05) correlations was yielded by the two series carried out at the same locale (Cl-C2: 7%), while highest similarities were those between Stas A and F (32% of the indices significant at P < 0.05). A more in-depth analysis based on curve-fitting by eye and cluster analysis (not shown), suggested several species groups with more or less clear preferences for a restricted depth interval. Figure 7 and Table 2 summarize these results, which are described as follows: Group I: species with abundance maxima near the surface: Subgroup 1: 0 m maximum and sharp decrease below. Subgroup 2: 0 m maximum and gradual decrease below. Subgroup 3: 25 m maximum. Subgroup 4: 50 m maximum and secondary 0 m maximum (Sta. Cl only). Subgroup 5: 50 m maximum. Subgroup 6: 0 and 100 m marked maxima. Group II: species with subsurface maxima: Subgroup 7: 100 m maximum. Group III: the deepest t&300 m species: Subgroup 8: 200 m maximum. Subgroup 9: 200 and 300 m maxima (one radiolarian only). Subgroup 10: 300 m maximum. Of these 38 radiolarians in the O-300 m data set, between 37 (Stas A and F) and 29 (Sta. Cl) were included in one or more groups, and none was excluded from all groups. Several profiles were not assigned to groups because of irregular or ambiguous vertical patterns. Similarities between the specific composition of groups I-III at the four stations were variable and generally rather low. Of the 34 species which were most abundant in the upper layers at one or more stations, only 14 entered group I at all four sites surveyed (asterisked in Table 2). Another seven entered group I at three of the four stations, and the remaining 12 only at two or one. Fifteen radiolarians had peak abundances at 100 m at one of the stations (group II), three at three stations, and only two taxa (Eucecryphalus sp. and Litharachnium tentorium) showed similar 100 m peaks at the four stations (Table 2). Finally, out of the 10 taxa which peaked at 200 or 300 m at one or more stations (group III), only Theocalyptra bicbrnis and Theocalyptra divisiana cornutoides showed coherent deep vertical patterns throughout the area (Table 2). Figure 8 summarizes similarities in radiolarians species compositions between stations for the three depth-intervals described on the basis of the vertical patterns defined. The uppermost layer is faunistically the most homogeneous throughout the area: on the average, ca 63% of the species which peak within this depth-interval are common to two or
Radiolarian
vertical
distribution
patterns
203
Fig. 7. Vertical distribution of the most abundant radiolarians in the O-300 m layer based on 2.5 m interpolated profiles. Shaded areas encompass all profiles grouped within each of the general patterns defined (see Table 2 for species included in each group).
more stations (asterisked in Table 2). Mid-depth and lower-depth taxa (groups II and III), however, vary considerably more between sites (26 and 30% in common, respectively), yet these similarities are still considerably higher than those between groups (9 to 4%) see Fig. 8). In addition to the abundant radiolarians, vertical ranges were also assessed (on the basis of maximum abundances throughout the water-column) for 28 taxa that were less abundant yet present at 23 of the four stations, and followed rather similar trends throughout the area. Table 2 illustrates the data in question: species 87 through seven (‘Upper 50 m” in Table 2) consistently peak between 0 and 50 m, and therefore have ranges similar to those described for group I. For taxa 43-127 (“O-100 m” in Table 2) and 139-52 (“25/50-100 m” in Table 2), peak abundances change from site to site between 0 and 100 m, with deepest maxima often occurring at Stas A and F. Radiolarians 129 through 28 (“100-300 m” in Table 2) center on 100-300 m. Finally, the last three taxa illustrated
204
S. A. KLING
and D. BOLTOVSKOY
Radiolarian ; . i i E i
E e : . E i : L
L .
L i
.
i
.
vertical
distribution
patterns
205
206
S. A. KLING
Percentage
and D. BOLTOVSKOY
of species
in common
3
Fig. 8. Similarities between radiolarian specific makeups for the &300 m vertical distribution groups I-III (cf. Fig. 7 and Table 2). Numbers denote species in common as a percentage of the total number of taxa encompassed by the two groups compared.
(ID nos. 9, 24 and 46, “200-300 + m” in Table 2) are consistently deep dwellers, with patterns similar to those of group III (Fig. 7). In agreement with the profiles for total shells (Fig. 3), the same radiolarian species tended to dwell shallowest in the water-column at Sta. F, and deepest at Stas Cl and C2. In the upper 300 m there did nt seem to be any relationship between preferred depth and differences in depth distribution from one station to another. In other words, most radiolarians occupied generally deeper strata at Cl-C2 than at A, and deeper at A than at F, regardless of their individual profiles. Thus, for the 13 consistently upper layer taxa (asterisked in Table 2), for example, the mean weighted population depth [WPD = C(ni . di)l%zi, where ni is the number of individuals at depth i] varied from 52 m at Sta. F, to 69 m at Sta. A, to S&91 m at Stas Cl-C2. For the rest of the 41 abundant radiolarians these values were 113, 137, 145 and 159 m, respectively (calculations based on O-300 m data only). The O-2000 m layer Thirty-nine of the 136 radiolarians identified had maximum abundance below 300 m at one or more stations (Table 3). Only three of these, however, were abundant (i.e. 220 individuals recorded in any one sample): Larcopyle buetschlii, Spongopyle osculosa and Spongurus sp. The latter two had fairly consistent patterns peaking generally at or below 300 m (Table 3, Fig. 9), but L. buetschlii’s abundances varied more between stations (Fig. 9).
Radiolarian
vertical
distribution
patterns
207
Table 3. Species with peak concentrations at depths of 300 m or more at one or more stations (selected from entire database). Data for each site are maximum abundance recorded (ind. mP3) and, in parentheses, corresponding depth. Frames highlight maxima at 300 m (thin line), and below 300 m (thick line). Last column gives maximum abundance in entire collection (ind. mP3). Taxa are sorted in descending order of maximum concentration recorded Results of ANOVA CZ/F 1 Cl/F
Total signif=ant
contrasts
1
16 1
1 Cl/C21
14 1
2 1
A/F
1 A/C2
18 1
1 A/Cl
10 1
12
S. A. KLINC and D. BOLTOVSKOY
Cyrtolagena
@
Larcopyle
buetschlii
Fig. 9 Vertical distribution
laguncula
Q
Spongopyle
osculosa
of deep-dwelling species. Numbers above profiles indicate scale maxima in ind. m-3.
Among the rare species, only Cyrtolagena laguncula and Sethoconus (?) sp. had maximum abundances below 300 m at the three sites with data from the deeper layers (Table 3, Fig. 9). In addition to the above, a few other radiolarians tended to peak below 300 m, yet their concentrations were too low to define reliable vertical trends: Botryostrobus aquilonaris, Siphocampe arachnea, Phormostichoartus corbula, Bathropyramis woodringi, Artostrobus annulatus and Sethoconus (?) sp. aff. S. (?) dogieli (cf. Table 3). Depth-abundance
relationships
The above-described depth preferences are probably to some extent related to overall abundances of the species in the entire database. Figure 10, which summarizes the stationto-station ranges in peak abundance for the 136 taxa, shows that deep-living forms are mostly located on the right-hand side of the graph (i.e. lowest mean abundances), whereas the upper-layer taxa are the most abundant. In addition, the station-to-station vertical ranges tend to be larger for the deeper, less abundant radiolarians. We suggest that this figure illustrates the combined consequences of several effects: (1) The uppermost layers host the numerically dominant species, whereas specific abundances decrease sharply at greater depths; (2) Vertical ranges tend to be better circumscribed in the upper layers, where environmental gradients with depth are steeper, than in the more homogenous meso- and bathypelagic layers. This trend has long been recognized, both for radiolarians (e.g. KLING, 1979), and for many other planktonic groups (VINOGRADOV, 1968); (3) Because deep-water species are less abundant, sample-size-related errors in the definition of their vertical ranges are larger. Thus, whereas (1) and (2) are actual relationships, (3) is an artifact.
Radiolarian vertical distribution patterns
Radiolarian
209
species
Fig. 10. Station-to-station ranges of depth of maximum concentration for all the radiolarians identified (hanging bars), as a function of their mean abundance in the entire collection (gray shading).
Comparison
with previous data
Although detailed comparisons of our results with previous studies is seriously restricted by methodological and nomenclatural differences, data summarized in Table 2 generally agree with, and amplify, previously published results. KLING'S (1979) survey of the vertical distribution patterns of polycystines of the central north Pacific is the study most closely comparable to the present one. Comparison of abundance maxima in the vertical profiles of 39 (out of total of 43 identified by KLING, 1979) radiolarians indicates that, with the exception of six taxa, vertical patterns derived from the present database encompass the ranges found by KLING (1979) in the central north Pacific. Only Cornutella profunda, Lamprocyrtis nigriniae, Theocalyptra bicornis, Botryostrobus aurituslaustralis, Dictyophimus infabricatus and Lophocorys polyacantha had deeper maxima in the central north Pacific than in any of the BMET stations. None of the above, however, exceeded 3.5% of all radiolarians in KLING'S (1979) collection or 3.7% in our database. Species-specific relative abundances, on the other hand, show differences consistent with warmer-water conditions at the INDOPAC sites (surface temperature ca 22-23”(Z), than at the sites covered in this work (see Fig. 2). Thus, in the O-100 m layer the typically warm-, oligotrophic-water radiolarians Heliodiscus asteriscus, Lithopera bacca, Phormospyris stabilis s.1. and Siphonosphaera polysiphonia were 5-165 times more abundant in KLING'S (1979) collection than off San Diego. Conversely, the BMET area O-100 m layer had higher proportions (5-50 times) of Botryostrobus aurituslaustralis, Pterocorys clausus, Spongocore cylindrica, Theocalyptra davisiana cornutoides and a few other characteristically colder water species. The BMET database is also generally consistent with the collection reported by BOLTOVSKOYand RIEDEL (1987) from O-100 m samples retrieved throughout 1972 in the California Current. Differences other than those attributable to variable sample-size and species-definition disagreements chiefly affect some of the deep-dwelling radiolarians which were not recorded in Boltovskoy and Riedel’s shallow tows (e.g. Artostrobus annulatus, Bathropyramis woodringi, Botryostrobus aquilonaris, Lamprocyrtis nigriniae, Cyrtolagena laguncula, Siphocampe arachnea, Spongopyle osculosa, and a few others; cf.
210
S. A. KLINC and D. BOLTOVSKOY
Table 2). Large dissimilarities in percentage contribution between the two collections were noticed for the deep-dwelling Theocalyptra davisiana s.1. and Spongocore cylindrica (ca lo-40 times more abundant at BMET), whereas the warmer-water Heliodiscus asteriscus, Polysolenia murrayana and Solenosphaera polyrnorpha were 8-13 times more abundant in BOLTOVSKOY and RIEDEL’S (1987) collection. Larcopyle buetschlii, however, whose maxima were at or below 100 m at all the sites surveyed (Appendices 1, 2), was approx. 20 times more abundant in the O-100 m California Current plankton than in the BMET samples. Hydrographic
control of radiolarian
patterns
Sparsity of contemporaneous hydrographic data for this study limits direct comparisons between species distributions and hydrographic patterns. Further complications arise from the complexity and instability of the California Current system, which lend importance to discrepancies in the time and spatial scale between hydrographic and plankton data collections. Nevertheless, some general patterns seem to emerge. As described above, analysis of geographic changes in radiolarian vertical distribution and assemblage makeup show that the two stations located at the extremes of the transect (A and F) are rather similar insofar as highest overall concentrations occur closest to the surface at these stations (Fig. 3), and assemblage compositions in the upper 25 m are more similar between these two locales, than between either one of them and Cl or C2 (Fig. 8). On the other hand, comparison of O-300 m relative abundances indicates that these two stations (F and A) differ most in their specific composition: of the 41 most abundant radiolarians, 21 showed significantly higher relative abundances at either Sta. F or Sta. A (Table 4 and Fig. 11). The nine species with highest values at Sta. F had, on the average, 5.1 ind. rnp3 at this site, and only slightly over 1 ind. rnp3 at the other three stations (Fig. 11). Six of these nine taxa are conspicuous members of group 1 (upper 50 m: cf. Tables 2 and 4). In contrast, species characteristic of Sta. A (12 in total, cf. Table 4) include several 100-300 m (Theocalyptra bicornis, Eucecryphalus sp, , Litharachnium tentoriurn, Dictyophimus infabricatus, D. gracilipes), and deep-water radiolarians (Spongurus sp.) (cf. Tables 2,3 and 4). In terms of the overall assemblages, differences between Stas A and F are quite significant insofar as they affect approximately half of the radiolarian shells recorded at Sta. F, and about 35% at Sta. A. This seemingly contradictory result is consistent with the regional circulation pattern as described above. We note that Central Water impinging on the western edge of the California Current, where Sta. F is located, is carried by the Southern California Eddy around Stas Cl and C2 and then north along the coast to the area of Sta. A (see “Oceanographic setting”). Since this Central Water is warmer than typical California Current waters, it overlies the latter near the coast. This may account for higher 0 m temperatures at Stas F and A. However, a thermally uniform 75 m upper mixed layer characterizes F, whereas temperatures drop sharply from the very top of the water-column at A (Fig. 2). This hydrographic pattern is consistent with radiolarian distributions. In the O-25 m layer of the area surveyed the 12 species which peak at the surface and drop sharply thereafter at Sta. A (group A-l in Table 2 and Fig. 7) comprise up to 11 times higher proportions of the assemblages at Stas F and A than at Cl and C2 (overall mean 3 times). Practically all these radiolarians are warm-water taxa, including well known indicators of
Radiolarian
vertical
distribution
211
patterns
Table 4. Variations in radiolarian relative abundances in the O-300 m layer of the four stations investigated. Body of the table indicates significant differences at P < 0.05 (one-way ANOVA and multiple contrasts according to Scheffi). Right-hand columns give the corresponding average percentages at each station (O-300 m only)
[14.77(300)1112.40 14 1Rhizoplegma 31
lheocatypha
boreale bicornk
0.52 (200)
0.34(200)
9.26(200)
0.65 (100)
WI “._. (200)
5.89
(100)
,I
5.89
1
p.12 500’ (
0.69 (0) 82 lLithe1iu.s sp. afi L. nautiloides
t t
135 45 24 54
16 21 117 20 89
165 134
1
Amphispyrti reticulota Potysolenia murrayana 1cornutelh profimda Heliodixus asteriscus ~Lamrocyclas maritalis $ichopilium bicorne 1Dictyophimus (?) sp. 1 Botryostrobus aquilonnrir 1Plagoniidae gen. et sp. indet. Lithopera bacca 1Callimitra cnro[ofae
0.: it 0.23 1 0.52 (lOa)
(4oo)l
1.83 (50)
~(1ooo)1.37(300)
11.72
11 (100)
(200)
IO.49 (300) 0.33 1.38 (0)
2mO1 1.37 (25)
1.38 (0)
0.52 (100)
91 1Siphocampe
--
98 1Phormostichoartus corbula 46 Bathropyramb woodringi 108 1Lithocampe sp. I
182 1Sethoconus
148 Lithe&u
(?) sp.
sp. 2
1 0.46 (50)
.-c-
0.
It- 0.06 (1000)
IO.09 ‘yoo’
1
1
M 1.38 1.37 0.77
arachnea
105 ~C)~tolagenalaguncu[a
1.72 1.55 1.38
212
S. A. KLING and D. BOLTOVSKOY
m
Station No
F (12
preference
spp.) (20
spp.)
0
c2 Stations
Cl
Fig. 11. Mean species-specific percentage contributions of the site-sensitive radiolarians to the G300 m assemblages of the four stations surveyed. Radiolarians are grouped according to their relative abundances at the four sites (cf. Table 4).
tropical-subtropical waters such as Didymocyrtis tetrathalamus, Eucyrtidium acuminaturn, Lipmanella dictyoceras, Pterocanium praetextum, Pterocanium trilobum, Pterocorys hertwigii, Sethophormis aurelia, Tetrapyle octacantha and Theocorythium trachelium. Although vertical profiles of these 12 radiolarians at A and F are very similar (Table 2)) a noteworthy difference is that their decrease with depth is much sharper at the coastal site (A), than at the oceanic one (F), in accordance with the difference in mixed-layer depths noted above. Thus, although at 0 m they comprise 32% (F) to 36% (A) of the fauna (unidentifieds excluded), at 50 m the percentage at F is 35%) and only 12% at A. In other words, similarities between F and A are high in the uppermost 25 m, but decrease rapidly with depth. This conclusion is clearly illustrated by results shown in Fig. 12. At 0 m Stas F and A are the most similar in their overall (relative) specific makeups. Below the surface, however, the oceanic Sta. F differs strongly from the other sites, and especially form Sta. A (Table 4). Some warm-water radiolarians, however, are abundant at the surface at Sta. F but show no substantial peaks at Sta. A. Among these, Cannobotryidae spp. et gen. indet., Spongosphaera streptacantha, Stylodictya spp. and Theopilium tricostatum, seem to be the most closely tied to site F; they account for 3.7% of total fauna at O-100 m at Sta. F, and only 0.6% at A. Pterocorys clausus and Theocorys veneris, also very abundant at the surface of Sta. F, have only very mild peaks at the surface of Sta. A. It is conceivable that all the above taxa are closely restricted to the Central Water and therefore cannot survive the trip to or conditions at Sta. A. Although the above interpretation is perhaps the most plausible, alternative hypotheses cannot be ruled out offhand. For example, the high abundances at Sta. F could represent a productivity event whose effects were not yet felt at Sta. A at the time of sampling. Comparison of our data with the pattern of polycystine distribution along the California Current derived by BOLTOVSKOY and RIEDEL (1987) reinforce the above conclusions. BOLTOVSKOY and RIEDEL (op. cit.), in a study of radiolarians from plankton tows collected
Radiolarian
vertical
distribution
patterns
213
Fig. 12. Similarities between overall (percentage) specific makeups at the four stations surveyed at 0, 50, 100 and 300 m. Clusters are based on MORISITA'S (1959) similarity index (HORN'S, 1966, modification), and UPGMA (SNEATH and SOKAL, 1973).
I
0.5
I
0.6
Fig. 13. Similarities between numbers for BOLTOVSKOY and California Current collected MORISITA'S (1959) similarity
I
/
1
I
0.7
0.8
0.9
1.0
radiolarian composition at 0 m at the stations surveyed and pooled RIEDEL'S (1987) station groups in O-LOO m plankton tows in the throughout 1972 (CC: California Current). Cluster is based on index (HORN'S, 1966, modification), and UPGMA (SNEATH and SOKAL, 1973).
throughout 1972 along the southern California Current grouped the assemblages retrieved into three distinct categories: colder-water associations restricted to the area north of 3234”N; transitional associations occurring between 25”N and 32-34”N; and warmer-water associations south of 25”N. Clustering of the above data (averaged numbers for pooled stations within each of the three categories), with 0 m results for our Stas (Fig. 13) shows
214
S. A.KLINC~~~D.BOLTOVSKOY
that sites F and A compare closely with the planktonic California Current warmer-water assemblage (“CC Warm” in Fig. 13), while Stas Cl and C2 cluster with the transitional assemblage of BOLTOVSKOYand RIEDEL (“CC Trans.“), and are also generally comparable to the colder-water one (“CC Cold”). The cold-water signal at subsurface layers of Sta. A may be reinforced by coastal upwelling, thus further enhancing the abundances of deeper-water radiolarians at this site. Such a mechanism is supported by the fact that, of the 11 radiolarians listed as “deep” (see “The O-2000 m layer” above), only two are present above 300 m at Sta. F (Spongopyle osculosa and Spongurus sp.), while at Sta. A nine of them occur in the same depthinterval. Similarly, some radiolarians that peak at subsurface depths at Stas F, C2 and Cl rise to a surface maximum at Sta. A (e.g. Dictyocoryneprofunda, Lithostrobus hexagonalis, and a few others, see Appendices 1,2). The inferred circulation responsible for radiolarian distribution patterns defined is schematically illustrated in Fig. 14. Implications
for the sedimentary record
Most distributional studies of Radiolaria based on surface sediment samples at some point dwell on comparisons between the bottom biogeographic imprint and hydrographic settings in the overlying mixed layer. These interpretations rely, either implicitly or explicitly, on the assumption that near-surface waters supply most or all of the shells recorded in bottom samples. Although the few direct comparisons of plankton and sedimentary information found important dissimilarities (PETRUSHEVSKAYA, 1971; RENZ, 1976; BOLTOVSKOY,1988, 1992; WELLING, 1990; SWANBERGand BJ~RKLUND, 1992),the latter were usually attributed to selective dissolution (PETRUSHEVSKAYA, 1981; RENZ, 1976; WELLING, 1990). However, a potentially important source of bias, or at least of blurring of differences between hydrographically dissimilar areas at the surface, is the flux of radiolarian shells from intermediate and deep layers. Because the physical environment in the meso- and bathypelagic realms is homogeneous throughout large areas of the World Ocean, its fauna is monotonous and it does not match biogeographic divisions in the upper waters. We attempted to compare the integrated absolute abundances of our shallow species, which are typically abundant over short depth intervals, with those of the deep radiolarians, which usually are less abundant over larger vertical ranges. Figure 15 illustrates the results of this analysis for Stas Cl and C2. Upper 50 m species (cf. Table 2) were assumed to yield live individuals between 0 and 100 m, while for the deep taxa (cf. Table 2) we averaged all their (presumably live) records between 300 and 2000 m. Although admittedly based on speculative assumptions, this figure suggests that depth-integrated abundances of deep-living radiolarians are higher than those of the shallow-water forms (approx. 4 times, according to data in Fig. 15). The integrated abundances are higher for the deep taxa (approx. 4 times at Sta. Cl, and about even at C2) even if calculated over 1000 rather than 2000 m. However, the contribution to sediment buildup from these deeper species, whose distributional patterns are not closely coupled with near-surface conditions, is dependent not only on their standing stocks, but also on their reproduction rates. It is conceivable that the nutrient-limiting conditions of meso- and bathypelagic depths render these sparse populations less productive than near-surface species. The degree to which these
Radiolarian
Fig.
14.
Inferred
circulation
vertical
affecting
distribution
the radiolarian
215
patterns
distribution
patterns
defined.
Radiolarian
cy 1.2 E 0-e nb 0.4 2 0.0
! ; r” % ‘pp~~qJ~ao:“‘““” r l-f-rrr
vertical
distribution
rrt-
0 ~ 0I- I-
Upper 50 m species at O-l 00 m Fig.
15.
217
patterns
N
Xr;E!
F 1?-
Deep species at 300-2000 m
Comparison of depth-integrated abundances of upper 50 m species, vs those of the deep species (cf. Table 2), at Stas Cl and C2. See Appendix 1 for species ID numbers.
presumably slower reproduction rates outweigh their higher absolute abundances remains an open question. The abundances of deep-water populations may be further enhanced by shells transported meridionally at depth from lower-latitude areas. As discussed above, several of the deeper-water radiolarians of our collection are absent or very scarce in O-100 m California Current waters throughout the year, but they are dominant components in north Pacific planktonic and sedimentary assemblages: Botryostrobus aquilonaris (ROBERTSON, 1975), Dictyophimus (?) sp. 1 (SACHS, 1973), Echinomma delicatulum (ROBERTSON, 1975), Heterucantha dentata (transitional species according to NIGRINI, 1970; SACHS, 1973; ROBERTSON, 1975), Rhizoplegma boreale (BLUEFORD, 1983), Siphocampe arachnea (KRUGLIKOVA, 1969; BLUEFORD, 1983), Spongurus sp. (TAKAHASHI, 1987, as Tholospyra group), Theocalyptra davisiana s.1. (cf. SACHS, 1973; BLUEFORD, 1983). These radiolarians seem to be abundant surface to subsurface dwellers closer to the pole but as the expatriated fractions of their populations move southward they sink deeper in the water-column and their densities decrease. This type of distribution pattern was demonstrated for some coldwater phaeodarian Radiolaria which submerge southward with the Intermediate subarctic waters (KLING, 1976), as well as for other radiolarians and zooplankton in general in all three oceans (VINOGRADOV, 1968; BOLTOVSKOY, 1992). BOLTOVSKOY (1988) reviewed mechanisms that can substantiate and enhance equatorward subsurface extensions of the ranges of cold-water planktonic organisms. An interesting example is Lithomitra arachnea, which is restricted to depths in excess of 200 m in our materials, peaking at 300-500 m (Sta. C2) to 1000 m (Stas Cl, F); and to lOO300 m in the central north Pacific (KLING, 1979). This species represents 25-60% of all radiolarians in surface sediments of the north Pacific north of 45-50”N, dropping to O-5% at the equator (KRUGLIKOVA, 1969). In sediments of the California Borderland it dominates the thanatocoenoses with contributions around 15-20% (KRUGLIKOVA, 1969; KLING, 1977). Yet in the O-100 m plankton it is absent altogether (BOLTOVSKOY and RIEDEL, 1987). The depths at which L. arachnea is found in extrapolar areas have low overall radiolarian standing stocks, and feeding conditions probably do not support very active growth and reproduction. Thus, we suggest that high abundances of this species in middle and low
218
S. A. KLING~~~
D.BOLTOVSKOY
latitude sediments are largely due to meridional transport, rather than to in situ production. Although vertical distribution profiles for L. aruchnea are not available for polar waters, the fact that it dominates high-latitude sedimentary assemblages implies high standing stocks of living populations which, therefore, are most probably associated with levels of maximum radiolarian abundance. In Arctic and Antarctic seas these maxima are located deeper than elsewhere, at about 20&400 m (TIBBS, 1967; BOLTOVSKOY and ALDER, 1992a,b; ALDER and BOLTOVSKOY, 1993). As mentioned in the introduction, the most widely used method in paleoenvironmental interpretations based on microfossil remains-the IMBRIE and KIPP (1971) factor analysis, transfer function technique-relies on the assumption that “core top faunas are systematically related to the physical nature of the overlying surface waters”. In this context, “systematically” means that the relationship in question has not varied in the time-span under investigation. However, the above-discussed contribution of laterally transported shells is dependent on the strength of intermediate and deep currents carrying-toward the equator-cooled waters that sink at the poles. Intensity of formation and output of these currents varied greatly between glacials and interglacials (BOYLE and KEIGWIN, 1987), conceivably affecting the contribution of cold-water polar shells to lower-latitude areas. Since meridional intermediate- and deep-water transport is strongest in the Atlantic and weakest in the Pacific, relationships between the sediments and the upper surface layer have probably been less uniform through time in the former than in the latter. Bathropyramis woodringi, Cornutella profunda, Cyrtolagena laguncula, and a few others, on the other hand, seem to be deep-water inhabitants in both low- and highlatitude areas. Their percentage contributions to overall sedimentary radiolarian assemblages do not increase towards the poles (NIGRINI, 1967; JOHNSON and NIGRINI, 1980,1982; BOLTOVSKOY, 1987). As far as paleoenvironmental interpretations are concerned, their records seem to be neutral background noise with no known information attached, other than the existence of abyssal ocean depths. REFERENCES V. A. and D. BOLTOVSKOY (1993) The ecology of larger microzooplankton in the Weddell-Scotia Confluence Area: horizontal and vertical distribution patterns. Journal of Marine l+arch, 51,323-344. BAKER A. DEC., M. R. CLARKE and M. J. HARRIS (1973) The N.I.O. combination net (RMT 1 + 8) and further developments of rectangular midwater trawls. Journal of the Marine Biological Association of the United Kingdom, 53, 167-184. BERNSTEIN T. (1934) Zooplankton Karskogo Morya po materialam ekspeditsii Arkticheskogo Instituta na “Sedove” 1930 goda i “Lomonosove” 1931 goda (Zooplankton des ntjrdlichen Teiles des Darischen Meeres). Trudy Arkticheskogo Instituta, Leningrad, 9, 3-58. BLUEFORD J. R. (1983) Distribution of Quaternary Radiolaria in the Navarin Basin geologic province, Bering Sea. Deep-Sea Research, 30,763-781. BOLTOVSKOY D. (1987) Sedimentary record of radiolarian biogeography in the equatorial to Antarctic western Pacific Ocean. Micropaleontology, 33,230-245. BOL~OVSKOY D. (1988) Equatorward sedimentary shadows of near-surface oceanographic patterns. Speculations in Science and Technology, 11,219-232. BOL~OVSKOY D. (1989) Las zonas de transici6n en la pelagial: biogeografia y paleobiogeografia. Mem. II Encontro Brasileiro de Plancton, Caiobd, 5-9 de dezembro de 1988, F. BRANDINI, editor, pp. 9-24. BOLTOVSKOY D. (1991) Holocene-upper Pleistocene radiolarian biogeography and paleoecology of the equatorial Pacific. Palaeogeography, Palaeoclimatology, Palaeoecology, 86,227-241. BOLTOVSKOY D. (1992) Current and productivity patterns in the equatorial Pacific across the Last Glacial Maximum based on radiolarian east-west and downcore fauna1 gradient. Micropaleontology, 38,397413. ALDER
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D. and V. A. ALDER (1992a) Paleoecological implications of radiolarian distribution and standing versus accumulation rates in the Weddell Sea. In: The Antarcticpaleoenvironment: a perspective on change, Antarctic Research Series, American Geophysical Union, 56, 377-384. BOLTOVSKOY D. and V. A. ALDER (1992b) Microzooplankton and tintinnid species-specific assemblage structures: patterns of distribution and year-to-year variations in the Weddell Sea (Antarctica). Journal of Plankton Research, 14, 1405-1423. BOLTOVSKOY D., V. A. ALDER and A. ABELMANN (1993a) Annual flux of radiolaria and other shelled plankters in the eastern equatorial Atlantic at 853 m: seasonal variations and polycystine species-specific responses. Deep-Sea Research I, 40, 1863-1895. BOLTOVSKOY D., V. A. ALDER and A. ABELMANN (1993b) Radiolarian sedimentary imprint in Atlantic equatorial sediments: comparison with the yearly flux at 853 m. Marine Micropaleontology, 23, l-12. BOLTOVSKOY D. and W. R. RIEDEL (1980) Polycystine Radiolaria from the Southwestern Atlantic Ocean plankton. Rev&a Espafiola de Micropaleontologia, 12,99-146. BOLTOVSKOY D. and W. R. RIEDEL (1987) Polycystine Radiolaria from the California Current region: seasonal and geographic patterns. Marine Micropaleontology, 12,655104. BOLTOVSKOY E. and H. A. LENA (1970) On the decomposition of the protoplasm and the sinking velocity of the planktonic foraminifers. Intationale Revue des Gesamten Hydrobiologie, 55,797-804. BOYLE E. A. and L. KEIGWIN (1987) North Atlantic thermohaline circulation during the past 20,000 years linked to high-latitude surface temperature. Nature, 330, 3540. BURCKLE L. H. (1981) Displaced Antarctic diatoms in the Almirante Passage. Marine Geology, 39, M39-M43. CARON D. A. and N. R. SWANBERG (1990) The ecology of planktonic sarcodines. Aquatic Science, 3, 147-180. CASEY R. E. (1966) A seasonal study on the distribution of polycystine radiolarians from waters overlying the Catalina Basin, Southern California. Ph.D. Thesis, University of Southern California, 136 pp. DWORETZKY B. A. and J. J. MORLEY (1987) Vertical distribution of Radiolaria in the eastern equatorial Atlantic: analysis of a multiple series of closely-spaced plankton tows. Marine Micropaleontology, 12, l-19. GOLL R. M. (1968) Classification and phylogeny of Trissocyclidae (Radiolaria) in the Pacific and Caribbean Basins. Journal of Paleontology, 42, 1409-1432. GOLL R. M. (1976) Morphological intergradation between Lophospyris and Phormospyris (Trissocyclidae, Radiolaria). Micropaleontology; 22, 379418. GOWING M. M. and S. L. COALE (1989) Fluxes of living radiolarians and their skeletons along a northeast Pacific transect from coastal upwelling to open ocean waters. Deep-Sea Research, 36,561-576. GOWING M. M. and D. L. GARRISON (1992) Abundance and feeding ecology of larger protozooplankton in the ice edge zone of the Weddell and Scotia Seas during the austral winter. Deep-Sea Research, 39,893-920. HAECKEL E. (1887) Report on the Radiolaria collected by H.M.S. Challenger during the years 1873-1876. Report on the Scientific Results of the Voyage of H.M.S. Challenger during the years 1873-1876, Zoology, 18, pp. i-clxxxviii + I-1803. HORN H. S. (1966) Measurement of “overlap” m comparative ecological studies. American Naturalist, 100, 419-424. IMBRIE J. and N. G. KIPP (1971) A new micropaleontological method for quantitative paleoclimatology: applications to a late Pleistocene Caribbean core. The Late Cenozoic Ages, Yale University Press, New Haven, pp. 71-181. JOHNSON D. A. and C. NIGRINI (1980) Radiolarian biogeography in surface sediments of the western Indian ocean. Marine Micropaleontology, 5, 111-152. JOHNSON D. A. and C. NIGRINI (1982) Radiolarian biogeography in surface sediments of the eastern Indian ocean. Marine Micropaleontology, ‘7, 237-281. JORGENSEN E. (1905) The protist plankton and the diatoms in bottom samples. Bergens Museum Skrifter, 1905, pp. 49-151, 195-225. KLING S. A. (1973) Radiolaria from the eastern north Pacific, Deep Sea Drilling Project, Leg 18. Initial Reports of the Deep-Sea Drilling Project, U.S. Government Printing Office, Washington, D.C., 18, 617-671. KLING S. A. (1976) Relation of radiolarian distribution to subsurface hydrography in the North Pacific. Deep-&a Research, 23, 1043-1058. KLING S. A. (1977) Local and regional imprints on radiolarian assemblages from California coastal basin sediments. Marine Micropaleontology, 2, 207-221. KLING S. A. (1979) Vertical distribution of polycystine radiolarians in the central North Pacific. Marine Micropaleontology, 4, 295-318. KRUGLIKOVA S. B. (1969) Radiolyarii v poverkhnostnom sloe osadkov severnoi poloviny Tikhogo Gkeana. In: BOLTOVSKOY
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Tikhii Okean, Mikroflora i mikrofauna v sovremennykh osadkakh Takhogo Okeana. Akademiya Nauk SSSR, Nauka, Moskva, pp. 48-72. MCMILLEN K. J. and R. E. CASEY (1978) Distribution of living polycystine radiolarians in the Gulf of Mexico and Caribbean Sea, and comparison with the sedimentary record. Marine Micropaleontology, 3, 121-145. MILLXMAN J. D. and K. TAKAHASHI (in press). Carbonate and opal production and accumulation in the ocean. In: GlobalsurJicialgeoJEuxes: Modern to Glacial, T. M. USSELMAN, W. HAY and M. MEYBECK, editors, National Academy Press. MORISITA M. (1959) Measuring of interspecific similarity and association between communities. Memoirs of the Faculty of Science, Kyushu University, Series E (Biology), 3,65-80. MORLEY J. J. and J. C. STEPIEN (1984) Siliceous microfauna in waters beneath Antarctic sea ice. Marine Ecology Progress Series, 19, 207-210. NICRINI C. A. (1967) Radiolaria in pelagic sediments from the Indian and Atlantic Oceans. Bulletin of the Scripps Institution of Oceanography, University of California, San Diego, 11, 125 pp. NIGRINI C. A. (1970) Radiolarian assemblages in the north Pacific and their application to a study of Quaternary sediments in core V20-130. Geological Society of America, Memoir 126, pp. 139-183. NIGRINI C. A. and T. C. MOORE (1979) A guide to Modern Radiolaria. Cushman Foundation for Foraminiferal Research, Special Publication 16, pp. Sl-S142 + Nl-N106. PELAEZ J. and J. A. MCGOWAN (1986) Phytoplankton pigment patterns in the California Current as determined by satellite. Limnology and Oceanography, 31,927-950. PETRUSHEVSKAYA M. G. (1967) Radiolyarii otryadov Spumellaria i Nassellaria Antarkticheskoi oblasti (po materiaiam Sovetskoi Antarkticheskoi Ekspeditzii). Issledovaniya Fauny Morei ZX(XII), Rezultaty Biologicheskikh Issledovanii Sovetskikh Antarkticheskikh Ekspeditsii 19.55-1958,3,5-186. PETRUSHEVSKAYA M. G. (1971) Spumellarian and nasselarian Radiolaria in the plankton and bottom sediments of the Central Pacific. In: The micropaleontology of oceans, B. M. FUNNELL and W. R. RIEDEL, editors, Cambridge University Press, Cambridge, pp. 309-318. PIELOU E. C. (1966) Shannon’s formula as a measure of species diversity: it’s use and misuse. American Naturalist, 100,463-465. POKRAS E. M. and B. MOLFINO (1986) Oceanographic control of diatom abundances and species distributions in surface sediments of the tropical and southeast Atlantic. Marine Micropaleontology, 10, 165-188. POPOFSKY A. (1908) Die Radiolarien des Antarktis (mit Ausnahme der Tripyleen). Deutsche Siidpolar Expedition 1901-1903,lO (Zool. 2), 3,183-305. POPOFSKY A. (1913) Die Nassellarien des Warmassergebietes. Deutsche Siidpolar Expedion 1901-1903,14 (Zool. 6) 217-416. REID J. L. (1965) Intermediate waters of the Pacific Ocean. The Johns Hopkins University Press, Baltimore, 85 PP. REID J. L. (1973) The shallow salinity minima of the Pacific Ocean. Deep-Sea Research, 20,51-68. REID J. L. G. I. RODEN and J. G. WYLLIE (1958) Studies of the California Current system. CalCOFI Progress Reports 7-l-56 to l-l-58, Marine Research Committee, California Department of Fish and Game, Sacramento, California, pp. 27-56. RENZ G. W. (1976) The distribution and ecology of Radiolaria in the Central Pacific; plankton and surface sediments. Bulletin of the Scripps Institution of Oceanography, 22, l-267. ROBERTSON J. H. (1975) Glacial to interglacial oceanographic changes in the northwest Pacific, including a continuous record of the last 400,000 years. Ph.D. Thesis, Columbia University, 355 pp. SACHS H. M. (1973) Quantitative radiolarian-based paleo-oceanography in late Pleistocene subarctic Pacific sediments. Ph.D. Thesis, Brown University, 201 pp. SHANNON C. E. and W. WEAVER (1949) Th e mathematical theory of communication. University of Illinois Press, Urbana, 125 pp. SNEATH P. H. A. and R. R. SOKAL (1973) Numerical taxonomy. Freeman, San Francisco, 573 pp. SWANBERG N. R. and K. R. BJ~RKLUND (1992) The radiolarian fauna of western Norwegian fjords: a multivariate comparison of the sediment and plankton assemblages. Micropaleontology, 38,57-74. TAKAHASHI K. (1983/84) Radiolaria: sinking population, standing stock, and production rate. Marine Micropaleontology, 8, 171-181. TAKAHASHI K. (1987) Radiolarian flux and seasonality: climatic and El Nido response in the Subarctic Pacific, 1982-1984. Global Biogeochemical Cycles, 1,213-231. TAKAHASHI K. and S. HONJO (1983) Radiolarian skeletons: size, weight, sinking speed, and residence time in tropical pelagic oceans. Deep-Sea Research, 30,543-568.
Radiolarian TIBBS J. F. (1967)
On some planktonic
protozoa
vertical taken
distribution from
the track
221
patterns of Drift
Station
ARLIS
I, 1960-1961.
Arctic
Institute of North America, 20, 247-254. VINOGRADOV M. E. (1968) Vertikalnoe raspredelenie okeanicheskogo zooplanktona. Nauka, L. A. (1990) Radiolarian microfauna in the northern California variability and implications for paleoceanographic reconstructions.
WELLING
Moskva, 320 pp. Current System: spatial and temporal M.Sc. Thesis, Oregon State University,
80 PP. J. G. (1966) Geostrophic Cooperative Oceanic Fisheries
WYLLIE
flow of the Investigations,
California Current at the surface and Atlas No 4, 13 pp, and 288 charts.
at 200 m. California
222
andD.
S. A. KLINC
APPENDIX
1.
BOLTOVSKOY
RADIOLARIAN
SPECIES
LIST
Species arc listed in alphabetical order, preceded by ID number. First set of square brackets gives maximum abundance recorded at each station (in ind. md3) and corresponding depth. Second set of square brackets furnishes a bibliographic identification reference and/or comments on the species’ taxonomic position. 91
Acunthodesmia sp. aff, A. vincufata (Mueller, Boltovskoy
and Riedel,
Giraffospyris
1987, Goll,
circumflsca,
41
Acunthodesmia
47
Acrosphuem spinosa (Haeckel,
1857)
as A. vinculutu
[F: 2.8-O;
Mueller.
C2: 0.5-50;
Open,
angular
Cl:
0.3-25;
structure,
A: 1.4-O]
similar
[?
to that of
19681
vincuhztu (Mueller,
1857)
[F: 1.4-O;
C2: ---; Cl:
---; A: 0.5-501
[Boltovskoy
and Riedel,
19871 Riedel, 181
Actinomma anturcticum (Haeckel, Rkdel,
68
132
[F: 0.1-750;
C2: 0.1-750;
Cl:
0.3-25;
A: ‘0.3-1001
C2: 3.7-50;
Cl:
20.0-o;
A: 2.8-O]
[Boltovskoy
and
1860)
[F: 5.5-O;
[Boltovskoy
and
19801
Actinomma leptodenum Echinomma
15
1860)
19801
(Joergensen,
leptodenmun
1900)
Joergensen;
[F: 0.5-50;
Boltovskoy
C2: 0.3-100;
and Riedel,
Cl:
---; A: ---I
[Kling,
1977,
as
19871
Actinomma sol Cleve, 1900 [F: 1.0-100; C2: ---; Cl: 0.5-50; A: 1.4-O] [Boltovskoy and Riedel, Actinommidae sp. et gen. indet. [F: ---; C2: ---; Cl: 0.1-1000; A: ---] [Various unidentified
19801
actinommids] 135
Amphiplectu acrostoma Haeckel, 1887 [F: ---; C2: 0. I-500; Cl: ---; A: ---1 [Petrushevskaya, Amphispyris reticuhzta (Ehrenberg, 1872) [F: 1.0-100; C2: 0.3-25; Cl: 0.2-500; A: 2.8-O]
77
Amphirhopdum
141
and Riedel,
19711 [Boltovskoy
19871
ypsilon Haeckel,
1887 [F:
1.4-O;
C2: ---; Cl:
0.2-200;
A: 1.4-O]
[Boltovskoy
and Riedel,
19871 170
Anthocyrtidium
40
Anthocyrtidium
1979;
84
Boltovskoy
same as specimens
somewhat
more
Amchnocomllium
with
C2: 0.3-25;
[F: 0.7-O;
Cl:
0.1-100;
a hyaline Cl:
0.2-50;
C2: ---; Cl:
0.1-400;
1908, except distally
coarser
Cl:
1987,
A: 0.2-3001 5.0-50;
under
A: 2.3-1001
the same name,
but
A: 2.3-501
[Probably
bears
same as Lithomelissa
to Psilomelissa
a slender
apical
spine,
tricuspidata
var.
and pores
on the
and irregular] Cl:
---; A: ---I [Similar
to but narrower
than
19711
1862 [F: 0.5-100;
C2: 1.4-50;
Cl:
0.3-25;
A: 0.3-1001
[Boltovskoy
19871
Amchnocorys (?) sp. aff. A. pentacantha Popofsky,
1913 [F: 4.8-O;
C2: 5.0-50;
Cl:
0.5-50;
19771
Artostrobus annul&us [Petrushevskaya,
[Kling,
peristome]
A: ---I [Similar
that the cephalis
sp. 2 [F: ---; C2: 0.2-300;
sp. in Pterushevskaya,
C2: 0.1-750;
and Riedel,
sp. 3 [F: 0.7-O;
Amchnoctwys circumtexta Haeckel,
[Kling, 185
1872)
19871
(?)
C2: 0.5-50;
by Boltovskoy
and Riedel,
become
Amchnocorullium
and Riedel, 169
illustrated
less campanulate,
of Popofsky,
thorax
11.7-O;
in Boltovskoy
Lophophaenoma 150
[F:
19871
sp. 1 [F: 2.8-O;
Amchnocomllium longer
ovate,
Haeckel
abdominalis 142
and Riedel,
1872)
zunguebaricum (Ehrenberg,
[Probably
thorucites 13
ophirense (Ehrenberg,
(Bailey, 19671
1856)
[F: 0.0-1000;
C2: 0.3-100;
Cl:
0.3-1ooO;
A: 0.2-3001
A: 1.4-O]
Radiolarian
46
Barhropyramis
20
[Kling, 19731 Borryosrrobus aquilonaris
woodringi
vertical
(Campbell
distribution
and Clark,
(Bailey,
1856)
1944)
223
patterns
[F: ---; C2: 0.5300;
[F: 0.1-1000;
C2: 1.4-300;
Cl:
Cl:
0.6-400;
1.7-200;
A: 0.5-3001
A: 0.5-3001
[Kling,
19791 9.5
Botryostrobus uurituduustrulis [Kling,
1979,
(Ehrenberg,
as B. au&us-ausrralis
carolotae Haeckel,
134
&&r&m
1887 [F:
17
? Culoeyclas monumentum Haeckel,
18
Cunnobotlyidae
1844)
[F: 6.2-100;
group;
Boltovskoy
1.4-25;
C2: 0.1-400;
C2:
15.5-50;
and Riedel, Cl:
Cl:
1987;
11.7-50;
A: 12.8-1001
Boltovskoy
---; A: 1.4-O]
and Vrba,
[Boltovskoy
19891
and Riedel,
19871 spp.
1887 [F: 0.5-100;
et gen. indet.
[F:
14.5-O;
C2: 0.3-100;
C2: 0.3-25;
Cl:
Cl:
2.1-50;
---; A: ---I [Haeckel,
18871
A: 0.5-501
[Various
unidentified
to Carpocanistrum
spp.
A and B in
catmobotryids] 119
Carpocanium sp. 1 [F: 4.1-O; Boltovskoy
118
and Riedel,
Curpocanium sp. 2 [F: 6.9-O; and B in Boltovskoy
4
C2: 0.9-50;
Cl:
0.9-200;
A: 1.6-1001
1987; with
tennina
teeth]
(Popofsky,
1913)
[Similar
[F: ---; C2: 0.1-500;
Carposphuem sp. aff. C. acaathophom (Popofsky, [Similar
to C. acanthophora
megaceros Hollande
Cladococcus [Boltovskoy
87
A: ---I [Similar
Cl:
to Carpocanistrum
0.1-400;
A: ---I
spp.
A
[Boltovskoy
and
19871
0.3-3001 56
2.4-25;
no teeth]
and Riedel,
Curposphaem ncunthophom Riedel,
60
C2: ---; Cl:
1987;
and Riedel,
Clathrocunium Riedel,
1980;
1913)
in Boltovskoy
[F: 0.7-O;
and Enjumet,
1960 [F: 4.6-50;
Kling,
as Cladococcus
courctutum Ehrenberg,
1977, 1860 [F:
C2: 0.2-300;
and Riedel,
8.3-O;
1987,
Cl:
but with
C2: 4.150;
1.2-200;
larger
Cl:
A:
pores]
1.7-200;
A: 4.1-5011
spp.]
C2: 1.0-25;
Cl:
0.7-O;
A: 12.4-O]
[Boltovskoy
and
19871
160
Collosphaem macropom Popofsky,
24
Cornutellaprofunda
1917 [F: 0.2-200;
C2: ---; Cl:
---; A: 8.3-O]
[Boltovskoy
and Riedel,
19871 Boltovskoy 151
123
[Somewhat
more
C. cervus;
includes
Corocdyptru
sp. aff.
83
cyrtohzgena 1979,
33
1873,
C. danaes
and Riedel,
group
[F:
than the category
Eucecryphalus
Kling,
Cl:
2.7-200;
A: 0.2-3001
[Kling,
1979;
craspedota Haeckel,
1.0-200;
illustrated
C2: 16.3-100;
Cl:
by Boltovskoy
(Joergensen)
1887 [F: 0.5-100;
22.4-50;
and Riedel,
of Kling,
A: 16.4-1001 1987,
under
the name
19791
C2: 0.3-300;
Cl:
0.7-25;
A: 2.7-501
19871 C2: 0.2-300;
Cl:
0.2-50;
A: 0.5-SO]
[Petmshevskaya,
19771
Cubotholus SP. [F: 0.1-1000; Cypussis irre&?daris Nigrini, 1977,
105
variable
C2: 0.5-300;
Cromyechinus borealis (Cleve, 1899) [F: 0.3-300; 1967;
71
1854 [F: 0.1-500;
19871
Corocalyptra cerws Ehrenberg,
[Boltovskoy 11
Ehrenberg,
and Riedel,
C2: ---; Cl: 1%8
0.1-750;
[F: 6.9-25;
A: ---I [Unidentified
C2: 0.3-25;
Cl:
Tholoniidae]
1.4-O;
A: 4.1-O)
Cl:
0.3-1000;
[Nigrini,
1968;
Klmg,
19791 S%Z~UIK.U~~ Haeckel,
as Stichopera
pecrinata
Dictyocephduspapi~losus
1887 [F: 0.1-1000; Haeckel
(Ehrenberg,
C2: 0.2-500;
[Kbg,
A: 0.3-1001
group] 1872)
[F:
1.0-300;
C2:
1.0-200;
Cl:
1.7-200;
A: 1.4-O]
[Nigini,
19671 79
Dictyocorynepmfbfda
102
Dictyophimus crisi~e Ehrenberg, Dictywhimus @wcif&es (Bailey,
Hymeniastrum 32
1977,
1979,
Ehrenberg,
euclidis;
1860 [F: 3.4-25;
Boltovskoy
as Psertdodictyophimus
and Riedel,
C2: 2.7-50;
[F: 25.9-100; gracilipes;
5.2-25;
A: 9.7-O]
[Kling,
1977,
19871
1854 [F: ---; C2: 0.1-750; 1856)
Cl:
Cl:
0.1-2000;
C2: 48.0-100; Boltovskoy
Cl:
and Riedel,
A: ---I 11.9-50; 19871
[Nig&i,
19671
A: 35.3-1001
[Khng,
as
224 146 103 8 117 130 55 69 10 131 104
152 78 36 171 139 136 110 54 43 48 59 1
66
S. A. KLING
and D. BOLTOVSKOY
Dic~ophimus infibticatus Nigrini, 1968 [F: 0.5-200; C2: 7.5-100; Cl: 1.0-200; A: 5.6-lOO] [Khng, 1977, 19791 Dictyophimus (?) kilhnati (Renz, 1976) [F: 0.9-300; C2: ---; Cl: 0.2-300; A: 0.7-lOO] [Renz, 1976, as Corocalyptra killmari] Dictyophbnus (?) sp. 3 [F: 0.7-O; C2: ---; Cl: 0.3-25; A: ---1 [Similar to Dictyophimus sp. aff. D. infabricatus Nigrini in Boltovskoy and Riedel, 19871 Dictyophimus (?) sp. 1 [F: 0.5-100; C2: ---; Cl: 1.7-500; A: ---I [Similar to Dictyophimus multispinus in Bernstein, 19341 Dictyophitnus (?) sp. 2 [F: 8.8-100; C2: 0.2-300; Cl: ---; A: ---] [Resembles Dictyophimus hirundo (Haeckel) group in Nigrini and Moore, 19791 Didymocyrtis tetmthnfamus (Haeckel, 1887) IF: 40.7-o; C2: 2.1-25; Cl: 18.5-25; A: 33. l-O] [Kling, 1979, as Ommatartus tetrathalamus; Boltovskoy and Riedel, 19871 Drynwsphaem (?) sp. [F: 0.7-O; C2: ---; Cl: ---; A: ---] [Provisional generic assignment according to Haeckel, 18871 Echinomma delicatulum (Dogiel, 1952) [F: ---; C2: 0.0-2000; Cl: 0.1-1000; A: ---I [Petrushevskaya, 1967; Khng, 19771 Echinommapopofskii Petrushevskaya, 1967 [F: 0.2-300; C2: 0.7-O; Cl: 1.4-O; A: 0.5-3001 [Khng, 19771 Enneaphonnis (?) sp. [F: 0.9-300; C2: 1.0-200; Cl: 0.7-O; A: 0.3-3001 [Generally similar to Enneaphonnis spp. in Petrushevskaya, 1971; probably conspecific with Sethophormis rot&a Haeckel in Renz, 19761 Eucecvphnlus sp. [F: 0.5-100; C2: 24.5-W; Cl: 30.6-50; A: 53.7-1001 [Boltovskoy and Riedel, 19871 Euchitonia eZeguns/furcuta (Ehrenberg, 1872) [F: 2.8-O; C2: ---; Cl: 0.7-25; A: 1.4-O] [Boltovskoy and Riedel, 19871 Eucyrtidium acuminatum (Ehrenberg, 1844) [F: 122.7-O; C2: 2.4-25; Cl: 8.6-25; A: 24.8-O] [Nigrini, 1967; Nigrini and Moore, 19791 Eucyrtidium hexagon&urn Haeckel, 1887 [F: 0.7-25; C2: ---; Cl: 0.325; A: ---I [Nigrini, 1967; Nigrini and Moore, 19791 EucyMium hexustichum (Haeckel, 1887) [F: 4.1-100; C2: 3.250; Cl: 3.1-25; A: 4.6-501 [Boltovskoy and Riedel, 19871 EucyrtiAum sp. [F: ---; C2: ---; Cl: 0.1-1000; A: ---I [Similar to E. acuminatum but narrower and with longer abdomen] Eucyrtidium sp. cf. E. culvertense Martin, 1904 [F: ---; C2: ---; Cl: ---; A: 0.2-3001 [Probably same as E. cienkowskii Haeckel in Boltovskoy and Riedel, 19801 Heliodiscus astetiscus Haeckel, 1887 [F: 2.1-O; C2: 0.1-400; Cl: 0.3-200; A: ---I [&Zing, 1979; Boltovskoy and Riedel, 19871 Helotholus histricosu Joergensen, 1905 [F: 2.1-O; C2: 0.9-50; Cl: 0.7-25; A: 0.7-1001 [Rling, 1977; Boltovskoy and Riedel, 19871 Hetemcantha dentutu Mast, 1910 [F: ---; C2: 0.1-750; Cl: ---; A: 0.3-lOO] [Nigrini, 1970; also similar to several other described taza, such as Cladococcuspinetum in Haeckel, 18871 Hexucontium laevigutum Haeckel, 1887 [F: 0.5-100; C2: 1.4-50; Cl: 0.3-1000; A: ---I [Boltovskoy and Riedel, 19871 Hexncontium sp. 1 [F: 4.1-100; C2: 6.2-300; Cl: 10.1-50; A: 5.5-501 [Kling, 1977, as H. pachydennum; large pores and conspicuous by-spines, similar to H. pachydennum in Joergensen, 1905, and to H. hostile Cleve in Boltovskoy and Riedel, 19801 Hexacontium sp. 2 [F: 0.5-300; C2: 2.3-50; Cl: 0.2-50; A: 0.5-501 [Probably same as H. entacanthum in Rling, 1977, and Boltovskoy and Riedel, 1980, 19871
Radiolarian
28
Lamprocyttis nigriniae (Caulet,
vertical
1971)
distribution
[F: 0.9-300;
225
patterns
C2: 2.3-100;
Cl:
3.3-200;
A: 1.3-3001
[Kbng,
1977,
19791 52 129
Lampromitmpambolica Popofsky, Lnmpromitm quadricuspis Haeckel, [Boltovskoy
16 72
and Riedel,
1913 [F: 0.5-100;
C2: 0.550;
1887 [F: 0.5-100;
C2:
Cl:
1.3-100;
---;
Cl:
A: 0.3-lOtI] 1.5-200;
[Popofsky,
19871
Lamprocycbzs maritalis Haeckel, 1887 [F: 0.3-1000; C2: 0.3-100; Cl: 2.1-200; A: ---I [Kling, L. m. polypora Nigrini] Larcopyfe buetschlii Dreyer, 1889 [F: 0.1-750; C2: 0.7-300; Cl: 9.3-200; A: 0.7-1001 [Kling, Boltovskoy
and Riedel
19131
A: 6.2-1001 1979,
as
1977;
19871
81
Larcopyle (2) sp. [F: ---; C2: 1.0-300; Cl: 0.5-300; A: 0.3-3001 [Conspicuously coarser external lattice than in Larcopyle buetschlii] Larcospim quadtanguta Haeckel, 1887 [F: 2.8-O; C2: 1.8-50; Cl: 6.2-25; A: 5.5-O] [Kling, 1977, 1979;
133
Lipmaneffa dictyocems (Haeckel,
9
Boltovskoy 1979;
and Riedel,
Boltovskoy
19871
and Riedel,
[F: 26.9-O;
as L. virchowii (Haeckel)]
C2: 9.1-50;
Cl:
7.6-25;
A: 40.0-O]
[Kling,
1977,
5
Lithamchnium
3
165
Lithelius minor Joergensen, 1900 [F: 2.1-25; C2: 1.6-100; Cl: 2.6-200; A: 13.8-O] [Kling, 1977, as L. minor (?)] Lithe&s sp. 1 [F: ---; C2: 0.2-300; Cl: ---; A: ---I [Unidentified Litheliidae, similar to Lithe&s sp. aff. L. alveolina Haeckel in Renz, 19761 Lithelius sp. 2 [F: ---; C2: 0.1-1000; Cl: 0.2-200; A: ---I [Unidentified Litheliidae; larger, coarser-meshed and shorter-spined than LitheZius sp. I] Lithelius sp. aff. L. nautiloides Popofsky, 1908 [F: 3.4-O; C2: 0.5-50; Cl: 0.4-1000; A: 1.4-O] [Probably conspecific with ?Pylospyra octopyle in Boltovskoy and Riedel, 19801 Lithocampe sp. 1 [F: 0.5-50; C2: ---; Cl: 0.1-1000; A: ---I [Nigrini, 1967, as Lithocumpe sp.] Lithomelissa setosa Joergensen, 1900 [F: 1.4-O; C2: 2.6-100; Cl: 11.7-25; A: 1.4-501 [Kling, 19771 Lithopem bacca Ehrenberg, 1872 [F: 0.7-O; C2: 0.1-750; Cl: 0.3-25; A: 1.4-O] [Kling, 1979; Boltovskoy
39
Lithostrobus haxagonalis Haeckel,
1979;
80 148 82 108 23
tentorium Haeckel,
1887) 1987,
Boltovskoy
and Riedel, Riedel, 154
1987;
and Riedel,
includes
C2: 5.2-100;
Cl:
7.1-50;
A: lO.l-1001
[Kling,
19871
many juvenile
specimens]
1887 [F: 0.5-100;
C2: 1.8-50;
1913 [F: 0.5-100;
C2: 0.2-300;
1887)
C2: 7.850;
Cl:
2.1-50;
A: 1.4-O]
[Boltovskoy
and
19871
Lophocoryspolyacantha Icling,
1860 [F: 3.1-100;
Popofsky,
Cl:
0.9-200;
A: ---I [Popofsky,
1913;
19791
168
Neosemantis distephanus (Haeckel,
92
Nephrospyris renifta Haeckel, 1887 [F: 1.4-O; C2: ---; Cl: ---; A: 0.3-1001 [Boltovskoy and Riedel, 19871 Porodiscus microporus (Stoehr, 1880) [F: 0.2-200; C2: 0.7-O; Cl: 1.4-O; A: 0.9-501 [Renq 19761 Peridium (?) SP. [F: 30.3-o; C2: 42.1-50; Cl: 49.1-25; A: 31.7-O] [Probably same as P. spinipes I-&&e1
Boltovskoy 74 85
in Casey, 70 184 38 50 98
and Riedel,
[F: 19.3-O;
Cl:
9.6-50;
A: 22.1-O]
[KJing,
1979;
19871
19661
Perypimmis circumtexta Haeckel, 1887 [F: Phormospytis s&bibs capoi Goll, 1976 [F: Phormospyris stabilis scuphipes GOB, 1976 Phormospyris stabilis stabifis (Goll, 1968) Phormostichoartus corbula (Harting, 1863)
---; C2: 0.2-300;
Cl:
---; C2: ---; Cl:
1.4-50;
[F: 0.5-100; [F: 2.1-O;
C2:
A: ---I
A: 3.2-501
1.4-50;
C2: 0.2-200;
[F: 0.1-1000;
0.2-300; Cl: Cl:
C2: 0.3-300;
2.5-50; 0.5-200; Cl:
19791
[Ming,
[GOB,
19761
A. 5.9-1001 A: 1.4-O]
0.6-1000;
[Go&
[KJQ,
19761 19791
A: 0.3~1001
[~hn~,
19791
89
Plagoniidae
gen. et sp. indet. [F: 1.6-100;
C2: 0.1-500;
Cl:
---; A: 0.3-1001
[Similar
to PI. 30, fig.
13 j,,
226 157 122 45 138
127 101 99 180 44 25 115 112 187 178 14 182
121 51 97 124 161 186 58 57
S.A. KLING andD. BOLTOVSKOY Popofsky, 19081 Plectauzntha (?) sp. aff. P. cremastoplegma Nigrini, 1968 [F: 17.9-O; C2: 0.2-300; Cl: 1.5-200; A: 5.5-O] [Provisional assignment according to Nigrini, 19681 Plegmosphaem sp. aff. P. lepticali Renz, 1976 [F: ---; C2: 0.9-300; Cl: 3.3-200; A: ---1 [Renz, 19761 Polysolenia murrayarm (Haeckel, 1887) [F: 2.7-25; C2: 0.0-1000; Cl: 0.1-500; A: 0.7-1001 [Boltovskoy and Riedel, 1987, as Acrosphaeru murrayuna] Psifomelissa (?) sp. [F: 19.3-O; C2: 7.8-50; Cl: 8.2-25; A: 2.8-O] [Probably related to Lophophaena sp. aff. L. apiculata Ehrenberg in Boltovskoy and Riedel, 1987; and to Psilomelissa galeata Ehrenberg in Popofsky, 19081 Pterocanium korotnevi (Dogiel, 1952) [F: 3.4-25; C2: 1.3-W; Cl: 2.1-O; A: 0.3-1001 [Nigrini and Moore, 19791 Pterocaniumpmetextum (Ehrenberg, 1872) [F: 36.4-25; C2: 0.5-50; Cl: 6.9-25; A: 40.0-O] [Boltovskoy and Riedel, 19871 Pterocanium sp. aff. P. gmndiporus Nigrini, 1968 [F: ---; C2: ---; Cl: 1.8-50; A: ---I [Similar to specimens illustrated by Nigrini, 1968, as P. gradiporus] Pterocanium sp. aff. P. trilobum (Haeckel, 1860) [F: 2.9-200; C2: ---; Cl: 1.025; A: 2.0-1001 [Atypical specimens resembling closely the named species] Pterocanium trilobum (Haeckel, 1860) [F: 37.2-O; C2: 2.4-25; Cl: 17.2-O; A: 26.2-O] [Kling, 19791 Pterocorys clausus (Popofsky, 1913) [F: 66.2-O; C2: 1.4-O; Cl: 2.1-O; A: 4.3-1001 [Klmg, 19791 Pterocorys hertwigii (Haeckel, 1887) [F: 41.4-O; C2: 4.1-25; Cl: 15.8-25; A: 63.5-O] [Boltovskoy and Riedel, 19871 Pterocorys minythomx (Nigrini, 1968) [F: 26.9-O; C2: 42.550; Cl: 50.1-25; A: 41.6-1001 [Kling, 1977, as Theoconus minythorax Nigrini; Boltovskoy and Riedel, 19871 Pterocorys sp. 1 [F: 1.8-50; C2: 6.5-25; Cl: 0.2-200; A: 1.6-1001 [Similar to P. macrocerus in Petrushevskaya, 19711 Pterocorys sp. 2 [F: 6.9-25; C2: 0.3-25; Cl: 0.3-25; A: 2.8-O] [Probably same as P. zancleus Mueller in Petrusbevskaya, 19711 Rhizoplegma boreale (Cleve, 1899) [F: 0.1-500; C2: ---; Cl: 0.2-200; A: 5.9-lOO] [Kling, 19771 Sethoconus (?) sp. [F: 0.1-1000; C2: 0.1-500; Cl: 0.3-1000; A: 0.3-1001 [Similar to Stichopilium variubile spinosum Popofsky in Petrushevskaya, 1967, but lacking postcephalic chamber divisions of Popofsky’s form] Sethoconus (?) sp. cf. S. (?) dogieli Petrushevskaya, 1967 [F: 0.1-500; C2: 0.2-200; Cl: 0.2-500; A: ---I. [Petrushevskaya, 19671 Sethophonnis aurelia Haeckel, 1887 [F: 71.7-O; C2: 3.7-50; Cl: 2.4-25; A: 46.9-O] [Boltovskoy and Riedel, 19871 Siphocampe arachnea (Ehrenberg, 1861) [F: 0.1-1000; C2: 0.2-300; Cl: 0.8-1000; A: ---I [Kling, 1979, as 5. aruchneu (Ehrenberg) group] Siphonosphaempolysiphonia Haeckel, 1887 [F: 3.4-25; C2: ---; Cl: 3.4-O; A: 5.5-O] [Kling, 1979; Boltovskoy and Riedel, 19871 Solenosphuempolymorpha (Haeckel, 1887) [F: 1.4-O; C2: ---; Cl: ---; A: ---I [Boltovskoy and Riedel, 19871 Spirocyrfis scalaris Haeckel, 1887 [F: 0.7-O; C2: 13.3-50; Cl: 5.3-50; A: 2.0-lOO] [Boltovskoy and Riedel, 1987, as S. scalariskomutella Haeckel (group ?)] Spongaster tetms Ehrenberg, 1860 irregdaris Nigrini, 1967 [F: 0.7-O; C2: 0.5-50; Cl: 4.1-25; A: 4.1-O] [Boltovskoy and Riedel, 19871 Spongocore cylitufriea Haeckel, 1860 [F: 8.8-100; C2: 1.4-O; Cl: 2.8-O; A: 7.8-501 [Kling, 1977, 1979,
Radiolarian
125
7
and Riedel,
227
patterns
C2: 8.6-300;
Cl:
8.9-25;
A: 4.3-1001
[Kling,
1979;
1987
Spongopyle osculosa Dreyer, 1889 [F: 3.0-400; C2: 2.4-300; Cl: 14.8-300; A: 12.4-3001 [Kling, 19771 Spongosphaera streptucantha Haeckel, 1862 [F: 11.7-O; C2: 2.7-50; Cl: 0.7-O; A: 0.9-501 [Boltovskoy and Riedel,
75
distribution
as S. puella; Boltovskoy and Riedel, 19871 Spongodiscus resurgens Ehrenberg, 1854 [F: 3.4-O; Boltovskoy
76
vertical
19871
Spongotrochus glacialis Popofsky,
1908 [F: 7.6-O;
C2: ---; Cl:
1.1-50;
A: 5.5-O]
[Boltovskoy
and Riedel,
19871 21
Sportgurus sp. [F: 1.5-500; C2: 2.9-500; Cl: 6.2-500; A: 5.5-O] [Kling, 1977, as Spongurus (?) sp.] Stichopilium bicorne Haeckel, 1887 [F: 0.2-400; C2: 1.8-50; Cl: 0.3-loo0, A: 0.3-1001 [Kling, 1979;
62
Stylacontatium bispiculum Popofsky,
27
Boltovskoy figs. 35
and Riedel,
19871 1912 [F: ---; C2: ---; Cl:
0.1-500;
A: ---I
[Skiing,
1973,
Plate 15,
1 l-141
Stylochlamydium venusturn Bailey, 1979;
Boltovskoy
and Riedel,
1856 [F: 0.9-50;
C2: 2.2-300;
Cl:
5.0-200;
A: 4.1-O]
[Wing,
1977,
19871
6
Stylodictya spp. [F: 11.0-o; C2: 2.2-200; Cl: 4.5-200; A: 3.9-1001 [Includes mainly forms similar to S. validispina in Kling, 19791 Tetrapyle octacantha Mueller, 1858 [F: 99.3-O; C2: 18.9-25; Cl: 37.8-25; A: 57.9-O] [Kling, 1979;
37
Theoculyptm bicornis (Popofsky,
73
Boltovskoy
and Riedel,
19871 1908)
[F: 0.5-200;
C2: 0.3-200;
Cl:
4.1-500;
A: 5.4-3003
[Kling,
1977,
19791 147
Theocalyptm aiwisiana (Ehrenberg, Cl:
106
29.9-200;
venen’s
Haeckel,
Boltovskoy
1967 [F: 0.9-300;
C2:
14.4-200;
19791 1862)
1887 [F: 41.4-O;
and Riedel,
[F: ---; C2: 0.7-200;
Cl:
0.5-100;
A: 2.9-1001
Boltovskoy
and Riedel
Tkeopilium tricostutum Haeckel, Riedel,
C2: 5.0-50;
Cl:
1.8-50;
A: 2.9-1001
[Rem,
1976;
W&g,
as Corocalyptra columba (Haeckel)]
1987,
Theocorythium tmchelium (Ehrenberg, 1979;
155
cornutoides Petrushevskaya,
1977,
19771
Theocovs 1979;
111
1862)
[Kling,
Theoculyptm davisiana davisiana (Ehrenberg, [Kling,
137
A: 3.1-3001
1872)
[F: 36.5-O;
C2: 0.5-50;
Cl:
4.8-25;
A: 15.2-O]
[a&
19871 1887 [F: 33.1-O;
C2: 25.6-50;
Cl:
2.1-200;
A: 2.3-501
[Boltovskoy
19871
90
zygocircusproductus
164
zygocircus sp. cf. 2. capulosus Popofsky,
and Riedel,
(Herhvig,
1879)
[F: 22.8-100;
C2: 1.4-50;
Cl:
1.5-200;
A: 5.9-1001
[Boltovskoy
19871
[Petrushevskaya,
1913 [F: 0.5-100;
C2: 0.2-300;
Cl:
0.2-200;
19711
163
Zygocircus (?) sp. [F: 4.1-O;
153
Unidentified
radiolarians
C2: 0.5-300; [F: 680.5-o;
Cl:
2.1-29;
C2: 241.2-100;
A: 1.4-O] Cl:
[D-shapedrings]
316.7-25;
A: 429.0-O]
A: 1.8-501
and
S.
A. KLING andD. BOLTOVSKOY APPENDIX
Radiolarian A 0
A 50
0.w 5.48 13.80 o.%? 0.00 0.00 57.94
0.00 0.w 0.00 0.00 0.00 0.00 1.38 1.38 0.043 0.04 0.W 0.00 0.00 0.00 0.00 2.78 5.52 0.04 1.38 1.38 4.14 24..%3 0.00 0.00 1.38 0.00 28.2, 0.W 0.03 0.00 1.38 0.00 0.00 33.1, 0.00
(ind. rnm3; top row gives station
counts
0.00 1.37
A loo 4.01 0.33 0.00 10.14
16.03 7.85 0.02 0.33 0.00 0.00 0.w 0.00 0.00 0.00 0.48 0.00 0.04 0.W 2.29 5.88 0.48 0.00 0.00 0.00 0.00 0.00 0.48 0.33 0.00 0.00 0.00 0.33 1.37 1.a, 0.00 0.W 4.25 0.W 0.33 0.02 0.00 0.04 .5.49 35.34 0.00 0.00 2.95 1.37 2.75 1.3, 0.00 0.33 3.66 5.80 0.02 0.65 0.02 2.29 0.40 0.W 0.00 0.65 1.37 7.85 0.W 0.65 0.00 0.04 0.00 0.33 0.00 0.33 0.00 2.29 0.00 0.04 3.86 4.12
A 300 2.28 3.43 0.00 2.12 4.24 0.16 0.w 0.33 0.00
0.16 0.00 0.33 0.16 0.00 0.00 0.00 0.40 0.00 0.10 0.10 0.00 3.10 1.31 4.4, 0.W 0.88 0.82 5.39 0.48 0.00 0.W 0.16 0.18 0.33 0.04 0.40 0.16 0.W
Cl 0 2.07 0.00 0.00 0.00 14.40 0.60 0.00 0.00 0.00
0.04 0.00 0.00 0.00 0.W 0.00 0.M) 0.00 0.00 2.07
0.W 2.07 1.38 0.00 4.83 0.00 4.83 2.78
0.00 0.00 0.04 0.00 0.00 0.00 17.24 0.00 0.W 0.00 0.00
7.78
4.14 0.00 0.00 0.04 0.00 0.00 4.14 0.w 0.00 0.W 5.52 4.14 1.38 1.38 %B8 0.09 5.52 1.38 0.W 1.38 31.73
-
0.48 0.00 0.00 0.00 0.48 0.00 0.00 0.00 0.00 O.MI 0.02 0.02 0.46 1.37
0.W 0.00 1.37
0.00 0.48 0.00 0.00 2.29 28.83
0.00 0.33 4.25 0.00 0.00 0.33 0.00 0.33 0.00 0.00 0.33 8.04
0.10 1.3 0.00 0.04 12.40 0.W 0.10 0.00 0.00 0.04 1.14 2.76 003 O.Oil 0.33 1.38 0.W 0.00 0.00 0.00 0.04 0.00 0.48 25.52
C-l 25
1.72
Cl 50
C-l loo
2
and depth, C-l 200
C-l 3co
left-hand C-l 400
column c-1 500
10.00 1.61 6.88 0.52 2.05 3.78 0.04 om 2.57 2.23 0.34 0.46 0.w 0.w 0.00 0.03 0.1, 0.00 7.09 0.6s 3.00 0.24 0.68 2.29 37.78 10.51 1.78 10.90 3.26 1.7, 1.37 0.34 0.23 0.00 0.17 0.17 0.23 0.23 0.34 0.00 0.08 0.17 0.00 0.w 0.00 0.w 0.00 0.00 0.34 0.52 0.00 0.11 0.00 o.cQ 0.00 0.00 0.00 0.00 0.00 0.17 0.00 0.23 0.1, 0.03 0.1, 0.1, 0.00 0.00 0.w 003 0.00 0.1, 0.w 0.00 0.00 0.00 0.17 0.02 0.00 0.00 0.00 0.46 0.53 0.17 0.00 0.00 0.00 0.00 0.23 0.00 2.08 0.00 0.00 0.23 0.00 0.W 0.00 0.03 0.00 0.00 0.00 0.69 2.06 0.00 0.17 0.17 0.M) 0.1, 0.6S 0.00 0.00 1.72 1.03 0.23 0.34 0.00 0.W 0.00 0.00 0.17 0.00 0.23 11.88 4.34 0.34 5.83 1.20 1.02 1.83 0.34 0.23 0.00 2.74 1.88 0.57 0.57 0.00 0.00 0.W 0.17 0.00 0.W 0.00 0.04 0.08 0.34 1.37 3.04 0.34 0.00 0.00 3.28 0.17 0.23 4.81 11.89 230 11.32 2.23 0.34 0.34 0.00 0.00 1.72 0.86 0.34 1.37 1.14 0.57 4.98 3.09 0.34 1.14 8.59 2.20 0.17 1.03 0.52 0.11 0.82 0.00 0.*3 0.03 1.03 0.00 0.57 4.12 0.34 2.51 0.17 0.5, 0.00 0.11 O.BS 0.00 2.08 0.W 0.88 0.W 0.23 0.46 0.00 5.03 0.48 0.00 0.00 0.00 0.03 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.89 0.00 0.08 0.5, 0.52 0.00 0.00 12.7, 10.29 1.08 2.57 0.00 0.1, 0.23 0.W O.Cil 0.06 0.00 0.00 0.00 0.11 0.00 0.00 0.00 0.17 0.34 0.57 0.34 0.34 0.00 0.M) 0.09 0.00 0.00 0.00 0.04 003 0.00 0.00 0.00 0.03 0.00 0.00 0.23 0.00 0.51 0.00 0.M) 0.W 2.40 080 0.1, 0.34 0.34 0.11 0.11 0.00 0.00 0.03 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.34 0.17 0.00 0.00 18.55 1.14 0.17 3.77 0.52 0.1, 0.11 0.29 1.72 0.34 0.23 0.48 0.68 0.9, 0.6S 2.06 0.1, 2.08 0.17 0.1, 0.48 4.12 0.46 003 0.89 0.34 0.00 0.00 0.W 0.23 0.09 0.00 0.17 0.23 0.23 0.00 0.00 0.00 1.20 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.1, 0.11 0.00 0.23 0.00 0.00 0.00 0.00 0.00 0.00 0.W 0.03 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.17 0.00 0.00 0.00 0.88 O.oB 0.5, 0.52 0.0-3 0.23 0.34 0.00 0.00 %?B 0.34 0.80 1.03 3.43 2.08 0.23 4.48 1.72 0.40 0.82 0.W 0.W 0.17 0.00 0.W 0.00 0.00 1.03 1.14 0.1, 0.34 0.34 0.02 0.1, 1.37 0.46 O.oB 7.03 ,477 8.42 0.24 0.00 0.00 0.00 0.17 0.00 0.00 0.00 0.68 0.00 0.00 0.04 0.00 0.04 0.W 5.15 0.9, 0.1, 1.20 0.34 0.11 0.48 0.00 0.00 0.04 0.00 0.00 0.00 003 8.18 0.00 0.1, 0.89 0.34 0.00 0.23 0.00 0.00 0.00 0.17 0.0.3 0.00 0.00 0.00 0.00 003 0.00 0.00 0.00 0.00 0.00 0.23 0.00 0.M) 0.M) 0.00 0.00 ‘lo.12 0x3 - 2.30 8.52 1.72 0.34 0.02 1.37
0.00 1.03
gives ID number)
C-l 750 0.57 0.00 0.00 0.18 0.40 0.00 0.00 0.18 0.08 0.W 0.00
0.W 0.00 0.65 0.00 0.00 0.18 0.08 0.00 0.24 0.04 0.4, 0.00 0.08 0.08 0.48 0.08 0.08 0.03 0.W 0.00 0.W 0.04 0.18 0.00 0.00 0.08 0.00 0.00 0.00 0.M) O.MI 0.33 0.33 0.08 0.00 0.16 0.00 0.00 0.00 0.00 0.00 0.W 0.W 0.24 0.24 0.00 0.00 0.04 0.18 0.00 0.04 0.04 0.08 0.00 0.4,
Cl moo
c-1 2wo
c-2 0
c-2 25
c-2 54
0.08 o.Bg o.00 1.83 0.00 0.00 0.00 0.00 0.00 0.w 0.00 0.00 0.15 0.w 0.34 0.00 0.20 5.54 18.88 ,417 0.09 0.00 0.34 2.74 0.00 0.00 0.w 0.00 0.03 0.00 0.00 0.00 0.08 0.00 0.00 0.00 0.00 0.17 0.00 0.M) 0.00 0.W 0.09 0.00 0.00 0.00 O.MI 0.W 0.03 0.00 0.00 0.00 0.00 0.00 0.00 0.00 1.37 0.04 0.M) 0.00 0.00 0.00 0.00 0.00 0.W 0.00 0.17 0.00 0.00 0.34 0.04 0.28 0.M) 0.00 0.00 0.00 0.34 0.03 0.00 0.00 1.83 0.00 0.03 2.08 1.7, 0.48 1.1, O.oB 0.00 0.00 0.00 0.28 0.00 1.38 1.03 0.0, 1.20 0.06 0.00 0.34 0.00 0.5, 0.03 0.M) 0.00 0.04 4.1, 0.60 1.03 5.94 0.00 0.09 0.00 0.00 0.W 2.57 0.12 0.69 2.06 ,.a7 0.00 0.00 0.69 2.40 0.9, 1.28 0.17 0.00 0.00 0.00 0.5, O.og 0.W 0.00 1.37 0.17 0.W 0.00 0.00 1.83 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.09 0.00 0.00 0.28 0.00 0.00 0.00 0.0, 0.77 0.00 0.88 2.40 1.37 0.00 0.W 0.00 0.W 0.34 0.08 0.00 0.00 0.00 0.00 0.03 0.W 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.W 0.00 0.W 0.00 0.04 0.00 0.88 1.03 3.88 0.00 0.M) 0.00 0.W 0.46 0.00 0.04 0.00 0.M) 0.86 0.03 1.38 2.Mi 1.37 0.43 0.W 0.80 2.40 4.1, 0.26 0.00 1.38 0.00 0.46 0.00 0.W 0.00 0.00 0.46 0.28 0.00 0.00 1.37 0.5, 0.08 0.00 0.00 0.00 0.00 0.00 0.00 0.W 0.00 0.00 0.W 2.29 0.W 0.00 0.00 0.03 0.00 0.00 0.00 0.00 0.00 0.00 0.M) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.34 0.00 0.88 O.oB 0.W 0.00 0.00 0.5, 0.17 0.00 0.W 0.00 0.77 0.00 0.00 0.00 0.00 0.00 0.M) 0.00 0.W 0.00 0.52 0.00 0.00 0.W 0.00 0.00 0.00 003 0.00 003 0.00 0.00 003 OS, 0.06 0.00 0.00 2.74 0.00 0.00 0.00 0.00 0.W 0.17 0.08 0.00 1.03 1.83 0.43 0.M) 0.00 0.00 0.48 0.00 0.00 0.M) 0.00 0.04 0.00 0.W 0.W 0.34 0.00 1.80 0.03 1.38 18.80 - 42.00 2.0, 0.00 0.w 0.86 1.54 0.08 0.w 0.5,
Radiolarian
A 0
A 50
A ,@I
12.42 0.00
1.83 0.00 4.58 0.00 0.M) 1,.44 0.00 0.00 0.00 2.28 0.W 0.w 0.04 0.00 0.00 0.00
1.3, 0.33 5.80 0.00 0.33 12.78
2.76 1.38 0.00 1.38 0.00 0.W 0.00 40.01 0.00 0.00 0.00 0.00 0.W 0.00 0.00 15.18 8.28 63.48 0.W 0.w 0.00 0.00 0.W 0.00 5.52 4.14 0.00 0.W 0.00
0.00 3.20 11.44 5.03 0.00 0.82 0.00 0.00 0.04 2.75 0.00 4.12 0.00 0.W 0.00
0.00 0.00 40.01
0.00 0.00 0.49
1.30 2.78 0.W 1.38 2.78
0.00 0.00 0.00 2.29 1.37
0.00 0.00
4.50 0.00
0.04 0.00 0.00 0.W 0.0-3 0.00 0.00 20.04 om 0.00 5.52
0.M) 0.02 0.w 0.00 0.w 2,Ol 0.48 226.90 0.00 2.22 1.83 0.00 0.00 0.48 1.83
8.28 0.09 1.38 0.00 1.38 22.07 1.38 0.00 0.00 2.76 0.00 2.76 0.00 0.00 0.00 - 0.04
0.48 l2.22 0.92 0.W 0.00 0.43 1.37 2.75 0.00 3.20 0.00 0.40 0.00
0.w 0.33 0.04 10.80 0.00 0.65 0.00 0.33 2.05 0.00 0.00 0.20 41.56 2.20 0.00 1.04 0.04 0.00 0.00 0.33 0.33 4.25 0.33 0.22 0.00 0.00 0.00 3.60 0.85 0.00 0.00 2.85 0.00 1.31 0.00 0.04 5.50 0.00 0.00 0.23 10.38 53.67 238.58 0.00 0.32 0.00
A 300 0.18 0.00 0.40 0.00 0.M) 2.12 0.03 0.00 0.W 0.08 0.00 0.09 0.33 0.00 0.00 0.00 0.18 0.33 4.90 1.14 0.00 0.33 0.00 0.00 0.00 0.33 0.10 0.10 0.10 0.49 0.04 0.40 0.00 0.40 0.00 0.18 0.00 0.00 0.00 0.33 0.00 0.00 0.42 3.10 0.00 0.0-3 0.98 1.3, 47.32 0.04 0.85 0.04
c-1
c,
0
25
0.09 0.00 0.w 0.00 0.00 1.38 0.M) 0.M) 0.00 4.83 0.00 0.M) 0.68 0.00 0.00 0.00 0.00 3.45 43.45 10.35 0.00
0.00 0.00 0.89 0.34 0.00 2.40 0.00 0.00 0.00 6.87 0.00 0.00 0.00 0.00 0.W 0.00 0.00 4.01 50.15 15.00
0.00 0.00 0.00 0.00 0.00 3.45 7.59 2.07 0.00 0.W 1.38 0.04 3.45 0.00 0.00 0.00 1.38 0.00 0.00 0.00 0.00 0.00 207 0.00 0.00 0.00 0.00 210.38 0.00 0.09 0.00
0.00 0.00 240 0.00 0.00 0.03 0.34 8.93 1.03 0.00 0.00
c-t 50
0.23 0.00 1.14 0.00 0.00 11.88 0.W 0.23 1.83 2.29 0.00 0.00 0.00 0.00 0.00 0.00 0.00 1.14 7.77 0.01 0.W 0.40 0.23 0.00 0.00 0.23 0.04 8.40 0.W 1.14 0.00 0.00
0.00 0.00 7.56
0.00 1.60
0.00 0.W 0.00 1.03 8.24
0.00 0.03 0.00 1.83 2.74
3.09 0.W 0.00 0.34 3.43 0.00 0.34 0.09 0.00 310.07 0.88
0.40 0.00
1.03 0.00
vertical
Cl lca 0.06 0.w 0.88 0.00 0.00 0.29 0.00 0.04 0.20 0.17 0.00 0.00 0.00 0.00 0.40 0.00 0.00 0.57 1.15 0.17 0.09 0.00 0.00 0.02 0.00 0.00 0.W 1.15 0.00 0.40 0.00 0.00 0.00 0.17 0.00 0.00 0.00 0.1, 0.11
distribution
c-1 200 0.00 0.00 1.54 0.00 0.00 11.15 0.34 0.00 0.17 1.54 0.W 0.17 0.24 0.17 0.34 0.W 0.W 0.89 10.47 1.54 0.00 0.86 0.00 0.M) 3.20 0.00 0.17 0.80 1.03 1.54 0.00 0.51 0.00 1.20 0.W 0.00 0.00 0.34 1.07
0.00 0.22 0.00 0.00 0.00 22.40 30.83 230.80 0.02
0.33 0.00 0.00 0.00
0.40 0.04 0.00 0.00 0.00 0.W
0.00 13.74 0.00 0.00 0.00 0.05 1.26 0.98 0.33 1.04 0.00 l.00 1.04
0.23 0.60 0.40 0.00 0.W 0.00 0.23 2.51 0.00 1.37 0.00 5.2* 0.00
c-1 300 0.w 0.m
2.58 0.17 0.00 0.00 0.W 0.00 0.17 0.17 0.00 0.02 0.00 0.00 0.00 2.58 0.00 0.00 0.24 0.w 0.00 0.86 0.00 0.W 0.W 0.w 0.04 0.W 0.17 0.00 1.55 0.00 0.00 0.00 0.00 0.17
42.83 0.00
1.43 0.00 0.00 0.00 0.00 0.06 0.75 0.W 0.40 0.03 0.34 0.00
2.40 0.34 0.w 0.W 0.M) 0.00 0.24 0.00 0.17 0.00 1.03 0.17
C-l 400 0.w
0.52 0.0.3 0.0.2
0.17 0.00 0.00 0.00 0.00 0.00 0.17 0.00 0.w 0.w 0.17 0.00
229
patterns
0.03 0.00 0.04 0.00 2.10 0.1, 0.11 0.M) 0.34 0.w 0.00 0.00 0.00 0.00 0.00 0.00 0.W 1.48 0.34 0.W 0.04 0.00 0.00 0.40 0.00 0.11 0.W 0.00 0.1, 0.00 0.00 0.00 0.23 0.00 0.00 O.MI
C-l 500 0.00 0.00 0.1, 0.00 0.00 3.86 0.11 0.57 0.M) 0.00 0.00 0.W 0.11 0.11 003 0.04 0.00 0.00 7.2, 0.23 1.72 0.W 0.W 0.22 1.w 0.11 0.w 0.04 0.00 0.1, 0.00 0.23 0.00 1.03 0.00 0.23 0.00 0.23
0.00 0.11 0.03 0.00 0.W 0.00 0.23 0.W 0.w 0.24 0.11 58.81 0.00
1.02 0.00 0.W 0.W 0.40 1.95 0.00 0.1, 1.26 1.14 78.83 0.00
0.57 0.W 0.00 0.00 0.23 0.00 0.00 0.00
0.22 0.11 0.00 0.00 0.23 0.11 0.00 0.34
0.00 0.00 0.00 0.00 0.00 0.00
0.04 0.00 0.00 0.00
0.00 0.23 0.W 0.02 0.03
0.11 0.00 0.34 0.00 0.43 0.11
Cl 750
o.00 0.00 0.00 0.00 0.00 0.24 0.08 0.08 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.W 0.w 0.00 0.04 0.00 0.00 0.00 0.00 0.03 0.00 0.00 0.00 0.02 0.W 0.16 0.04 0.00 0.04 0.08 0.00 0.00 0.08 0.00 0.W 0.04 0.08 0.W 7.00 0.00 0.08 0.03 0.00 0.03 0.00 0.00 0.W 0.W 0.00 0.00 0.00 0.00 0.00 0.60 0.00 0.00 0.00 0.00 0.00
Cl lK!o 0.w 0.w 0.03 0.00 0.04 2.44 0.77 0.80 0.00 0.09 0.M 0.00 0.17 0.34 0.17 0.00 0.00 0.03 2.0, 0.00 0.00 0.00 0.17 0.00
Cl *cm 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.03 0.00 0.00 0.08 0.03 O.MI 0.20 0.00 0.04 0.00 0.12 0.00
c-2
0
25
0.w 0.0-3 0.00 0.W 0.04 O.OS 003 0.00 0.00 0.00 0.M) 0.00 0.00 0.00 0.04 0.04 0.04 0.00 4.85 0.88
0.00 0.00 0.28 0.09 0.51 0.00 0.00 0.00
0.00 0.00 0.00 0.w 0.00 0.W
0.00 0.00 0.00 0.00 0.00 0.00 0.00 4.15 0.09 0.00
0.00 0.12 0.00 0.03 0.00 0.00 0.00
0.00 0.89 0.00 0.80 0.W 0.W 0.W
0.28 0.00 0.00 0.00 0.00 0.00
0.04 0.03 0.00 0.00 0.00 0.00
1.71 0.17 0.00 0.89 1.54 OS.43 0.17 0.77 0.00
0.44 0.00 0.06 0.06 0.00 3.52 0.00 0.00 0.00
0.W 0.00 0.00 0.00 0.00 0.00 2.08
0.00 0.00 0.00 0.00
0.00 0.W 0.00 0.00
0.00 0.43 0.00 0.53 0.00 0.00 0.00 o.*o 0.34 0.09 0.34 0.00
0.00 0.00 0.00 0.00 0.00 0.00 0.w 0.03 0.W 0.00 0.03 0.03
0.00 0.00 0.00 0.04 0.00 0.00 0.w 0.00 0.W 0.043 0.00
0.00
0.02
0.W
0.17 0.00 0.09 0.W 0.20
0.00 0.00
c-2
0.00
0.00 0.w 0.00 0.00 72.00 0.00 0.00 0.00 0.00 0.W 0.00 0.00
1.03 0.00 0.88 0.34 0.0.2 0.W 0.00
C-2 50 0.w 0.w 1.37 0.40 0.00 15.04
0.00 0.00 0.00 0.00 0.00 0.04 ,5.02 4.1,
0.00 0.00 0.W 0.48 0.00 0.05 0.00 0.00 0.00 0.00 0.00 0.40 42.51 2.28
0.W 0.W 0.00 0.00
0.m 0.0, 0.00 0.W
0.00 0.00 0.00 5.42 0.M)
0.w 0.00 O.Ml 8.86 0.w
0.W 0.00 0.00 0.00 3.09 0.00 0.34 0.03
0.W 0.00 0.05 0.00 0.14 0.00 0.00
0.03 0.00 0.W 0.W 0.00
3.42 5.14 O.BO 0.W 0.00 0.00 0.00 0.00 0.00 0.24 0.00 129.80 0.00 0.34 0.00 0.00 0.00 0.00 0.W 0.00 3.08 2.74 0.00 0.00 0.34 0.00 1.37 0.w 0.0-3 0.M) 1.37 0.5,
0.00 5.03 7.77 3.x1 0.00 0.00 0.00 0.00 0.w 1.37 16.00 0.W 149.03 0.00 0.M) 0.00 0.00 0.00 0.00 7.77 5.03 0.40 0.M) 0.00 3.60 0.00 0.00 0.00 13.28 1.03
230
7 3 4 5 6 7 8 8 10 11 13 14 15
16 17 18 20 21 23 24 25 27 28 32 33 35 36 37 38 39 40 41 43 4-I 45 40 47 48 50 51 52
54 55 Ea 57
5% 59 80 82 86 08 Es 70 71 72 73 74 75 78 7, 78 79 90 B1 82 83 a4 E
S. A. KLING
and D. BOLTOVSKOY
c-2 100
C-2 200
CL? 300
c-2 400
c-2 503
G2 750
c-2 1003
c-2 2ooo
F 0
F 25
F 50
0.w 1.63 0.00
1.03 0.03 0.03 0.86 2.40 0.08 0.00 0.00 0.W 0.00 0.00 0.00 0.00 0.1, 0.00 0.00 0.68 0.1,
6.17 0.34 0.00 1.71
0.78 0.00 0.043 0.76 1.53 0.22 0.W 0.11 0.00 0.00 0.W 0.00 0.00 0.00 0.W 0.00 0.11 0.33 0.W 0.22 0.00 1.0s 0.55 1.m 0.03 0.33 0.11 0.22 0.11 0.11 0.W 0.00 0.22 0.11 0.00 0.00 0.00 0.00 0.00 0.1, 0.00 0.11 0.11 0.44 0.11 0.11 0.W 0.00 0.00 0.00 0.00 0.00 0.11 0.1, 0.22 0.44 0.W 0.00 0.11 0.W 0.00 0.33 0.00 0.00 0.00 0.00 0.00 0.55
1.04 0.12 0.12 0.35 1.05 0.00 0.00 0.12 0.00 0.00 0.00 0.00 0.00 0.00 0.04 0.00 0.23 0.23 0.12 0.48 0.00 2.90 0.23 OH 0.12 0.58 0.12 0.00 0.23 0.12 0.00 0.00 0.12 0.M 0.00 0.03 0.03 0.W 0.12 0.23 0.00 0.00 0.98 0.35 0.00 0.00 0.12 0.00 0.M 0.00 0.00 0.00
0.28 0.0, 0.00 0.00 1.m 0.0, 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.14 0.00 0.W 0.00 0.07 0.14 0.0, 0.0, 0.83 0.1, O.sB 0.00 0.83 0.28 0.28 0.00
0.30 0.04 0.00 0.23 0.76 0.04 0.00 0.00
0.07 0.00 0.00 0.2, 0.29 0.03 0.W 0.W 0.02 0.00 0.W 0.00 0.00 0.07 0.00 0.00 0.02 0.00 0.00
3.45 0.00 0.00 0.00 88.29 11.72 0.60 0.00 0.00 0.00 0.00 0.00 0.69 0.00 0.00 14.48 0.00 0.00 I.38 0.W 88.18 1.38 0.00 13.,0 0.00 0.00 122.73 0.00 0.04 0.W 0.59 1.38 2.0, 37.23 0.00 0.00 0.03 0.W 2.07 71.7, 0.00 2.0, 40.68 2.0, 0.00 0.W 0.00 0.69 0.00 0.00 0.00 0.88 0.00 2.0, 0.00 11.03 0.00
2.00 2.00 0.00 0.00 62.48 *.@I 0.00 0.00 0.00 0.00
5.23
3.92 0.00 0.w 0.00 0.W 0.00 0.00 0.00 0.00 0.33 0.33 0.00 0.W 0.08 2.61 0.00 0.33 0.W 2.20 48.04 0.00 0.08 0.00 0.04 1.31 0.08 0.W 0.00 0.33 1.88 0.00 0.03 0.00 0.04 0.00 0.65
0.00 0.W 0.08 1.31 0.33 0.W 0.33 0.00 0.00 0.00 0.33 0.00 0.00 0.33 0.00 ,.$-a 0.03 0.00 0.33 0.00 0.00 0.95 0.00 0.00 0.23 0.00 0.33 10.78
2.40 0.51
0.00 0.1, 1.03 5.13 1.03 0.5, 0.00 0.34 0.5, 0.24 0.00 0.00 0.1, 0.08 0.00 0.17
0.00 0.00 0.17 0.00 0.00 0.00 0.17 0.51 0.51 0.04 0.W 0.00 0.00 0.34 0.04 0.00 0.1, 0.00 a.@3 2.22 0.00 0.00 1.7, 0.M 0.00 0.1, 0.17 0.17 0.00 0.00 0.M 1.e3
5.48
0.00 1.03 0.1, 0.00 0.00 0.1, 0.00 0.1, 1.3, 0.51 0.51 0.5, 0.04 0.51
0.60 5.88 0.51
2.23 0.24 0.00 0.34 0.51
0.00 0.00 0.09 1.03 0.W 0.51 0.00 0.00 0.04 0.00 0.00 0.00 0.88 0.34 0.03 0.1, 0.17 0.W 0.00 0.00 0.17 0.1, 0.@8 1.20 0.00 0.00 2.40 0.W 0.00 0.51
0.17 0.34 0.M 0.W 0.17 3.28
0.00 0.23
0.04 0.58 030 0.00 2.20 0.00 0.00 0.12 0.00 0.11 0.00 0.00 0.00 0.93
0.0, 0.0, 0.00
0.W 0.21 0.00 0.0, 0.0, 0.0, 0.w 0.00 0.00 0.0, 0.42 0.21 0.00 0.14 0.00 0.04 0.w 0.W 0.00 0.00 0.04 0.14 0.00 0.21
0.00 0.00 I.40 0.00 0.09 0.14 0.0, 0.00 0.07 0.00 0.14 0.0,
0.04 0.00 0.00 0.04 0.0-l 0.04 0.00 0.00 0.w 003 0.08 0.04 0.04 0.11 0.08 0.00 0.23 0.W 0.04 0.08 0.00 0.00 0.00 0.00 0.04 0.04 0.00 0.00 0.00 0.00 0.04 0.W 0.00 0.19 0.04 0.04 0.03 0.00 0.08 0.00 0.00 0.00 0.00 0.04 0.08 0.00 0.04 0.M 0.00 0.48 0.00 0.00 0.11 0.00 0.04 0.04 0.04 0.0-2 0.23
0.15
0.00 0.12 0.02 0.38 0.05 0.34 0.00 0.0, 0.w 0.05 0.00 0.w 0.05 0.07 0.00 0.02 0.00 0.W 0.00 0.00 0.00 0.W 0.07 0.02 0.05 0.00 0.00 0.05 0.00 0.00 0.00 0.00 0.02 0.05 0.02 0.28 0.00 0.00 0.0, 0.M 0.00 0.05 0.00 0.W 0.00 0.00 0.00 0.05
0.00 0.00 0.00 0.w 0.00 2.08 0.03 0.00 0.0s 0.00 40.51
0.04 0.00 6.8, 0.w 0.69 70.03 0.00 0.00 0.03 0.00 0.00 0.00 35.01 2.75 0.02 0.00 0.00 0.00 58.04 0.00 0.00 24.72 1.3, 0.69 0.00 0.00 0.w 0.00 0.00 0.00 0.00 0.00 8.57 0.09 2.75 0.00 4.12 0.04 0.W 2.06 3.43 0.00 0.00 2.78 1.37 3.45 2.08 0.00 0.w 2.78 2.08 30.34 8.18
F 100
F .mo
1.37 0.00 0.00 0.00 35.18
4.14 0.00 0.00 3.11 23.82
0.86 0.w 0.w 0.52 13.57
8.23 0.00 0.00 0.00 0.00 0.w 0.00 0.w 0.00 0.00
1.04 0.52 0.00 0.w 0.w 0.00 0.00 1.04 0.00 0.52 0.52
0.00 0.00 0.00 0.04 0.00 0.00 0.00
0.17 0.00 0.w 0.00
0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 2.58 0.W 0.00 1.03 0.00 0.00 0.52 0.52 0.W 0.34 0.1, 0.00 0.00 2.58 0.00 0.00 0.W 0.00 0.00 1.03 0.04 0.00 1.72 0.17 0.88 0.00 0.00 0.03 0.00 0.34 0.00 0.04 0.00 0.00 0.00 0.34 0.1, 0.00 0.00 0.00 0.04 0.17 0.00 0.5S 0.1, 0.00 0.W 1.03
0.00 0.00 0.W OS-XI 0.00 0.17 0.00 0.00 3.00 0.5, 0.86 1.88 1.03 0.00 0.60 0.1, 0.00 0.00 0.00 0.00 0.00 0.88 0.00 0.00 0.00 0.00 0.00 2.5, 0.00 0.00 0.68 0.1, 0.34 0.00 0.1, 0.34 0.W
2.74 0.00 0.00 0.03 0.00 21.02
0.48 0.00 2.23 0.00 0.01 39.30 0.00 0.00 0.40 0.00 0.00 0.w 15.08 0.00 0.04 0.03 0.03 0.00 39.30 0.W 0.91 15.54 4.5, 0.46 0.00 0.04 0.00 0.00 0.04 0.40 0.02 0.00 133 0.00 1.3, 0.00 0.00 0.00 0.47 0.91 1.83 0.00 2.29 1.M 0.00 0.00 1.37
0.00 0.00 0.52 0.00 24.88
0.w 0.W 25.80 0.04 0.W 6.73 0.04 0.52 0.52 0.52 0.00 0.52 5.70 0.04 0.00 0.00 0.00 0.W 8.80 0.52
0.03 3.11 1.04 8.80 0.00 0.52
0.00 0.00 0.00 0.00 0.00 0.02 0.00 0.00 2.58 0.00 1.04 0.52 0.W 1.04 2.0, 0.00 0.00 0.00 0.00 0.00 13.88
F 3w 1.37 0.w 0.00 0.00 0.77
0.34 0.00 0.00
0.51
0.00 0.04 0.00 0.04 0.W 1.37 0.00 0.17 1.37 0.1, 0.1, 0.1, 0.04 0.51 0.17 0.W 0.04 1.03
F 400
F 500
F 750
F loco
0.23 0.48 0.00 0.23 5.76 0.12 0.w 0.00 0.00 0.00
0.34 0.11
0.00 0.03 0.00 0.03 l.w 0.00 0.00 0.00 0.04 0.00 0.00 0.00 0.00 0.00 0.00
0.w 0.0s 0.00 0.23 1.57 0.00 0.03 0.00 0.w 0.0-3 0.w 0.00 0.00 0.20 0.00
0.00 0.03 0.00 0.00 0.00 0.03
0.06 0.06 0.00 0.00 0.00 0.11
0.W 0.00 0.00 0.W 0.00 0.04 0.00 0.23 0.00 0.00 0.81 0.48 0.35 1.50 0.w 0.23 0.58 0.00 0.00 0.12 0.03 0.00 0.00 0.35 0.00 0.00 0.00 0.00 0.00 0.02 0.00 0.00 0.00 0.35 0.35 0.12 0.00 0.12 0.00 0.00 0.00 0.W 0.00 0.23 0.12 0.58 0.12 0.12 2.89 0.00 0.12 0.00 0.00 0.04 0.00 0.04 0.00 0.23
0.48 3.30 0.00 0.00 0.00 0.00 0.00 0.00 0.11 0.00 0.17 0.00 0.00 0.00 0.W 0.57 1.48 0.11 0.34 0.00 0.00 0.23 0.17 0.11 0.1, 0.00 0.04 0.w 0.11 0.00 0.00 0.00 0.00 0.5, 0.W 0.00 0.40 0.23 0.11 0.00 0.08 0.00 0.00 0X-J 0.W 0.00 0.W 0.00 1.08 0.W 0.21) 0.00 0.00 0.00 0.00 0.06 0.08
0.23 _s 0.26 0.08 0.14 0.20 0.86 0.00 0.14 0.00 0.W 0.00 0.23 0.00 0.1, 0.08 0.W 0.00 0.00 0.W 0.W 0.00 002 0.00 0.03 0.00 0.17 0.00 0.04 0.00 0.00 0.08 0.09 0.00 0.00 0.03 0.00 0.11 0.20 0.00 0.00 0.00 0.00 0.1, 0.34 0.00 0.03 0.00 0.03 0.00 0.00 0.W 0.00 0.00 0.03 0.00 0.W 0.03 0.W 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.14 0.03 0.00 0.23 0.00 0.00 0.00 0.00 0.1, I.04 0.00 0.03 0.W 0.03 0.03 0.1, 0.00 0.00 0.08 0.03 0.00 0.M 0.00 0.08 0.00 0.03 0.09 0.09
Radiolarian
vertical
distribution
231
patterns
c-2
c-2
c-2
c-2
c-2
c-2
c-2
c-2
F
F
F
100
m
300
4m
Em
750
loo0
2Mx)
0
25
53
0.00 0.51 0.w 0.00
0.w 1.03 0.00 0.00
F too
F 200
F 3w
F 400
F 5ca
F 750
F 1oocl
0.w 0.00
0.w
0.03 0.00 0.w
8.57 0.34
0.87 0.11 0.w 0.00 0.00 0.00 0.44
a00 2.07
0.34 0.00
0.00 0.08
0.00 cLcK7
0.w CKO
0.w 0.00
0.w 0.00 0.51 0.17 0.w 0.17
0.17 0.00
0.00 0.00 0.17 0.00
0.22 0.00 0.00 0.00
0.w 0.06
0.w cl.03 0.00 0.44
0.w 0.23
0.w 0.00 mm 0.14