A. BUZAS Microdistribution of Foraminifera in a Single Bed of the Monterey Formation, Monterey County, California SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION Emphasis upon publication as a means of “diffusing knowledge" was expressed by the first Secretary of the Smithsonian. In his formal plan for the Institution, Joseph Henry outlined a program that included the following statement: “It is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge." 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The publications are distributed by mailing lists to libraries, universities, and similar institutions throughout the world. Papers or monographs submitted for series publication are received by the Smithsonian Institution Press, subject to its own review for format and style, only through departments of the various Smithsonian museums or bureaux, where the manuscripts are given substantive review. Press requirements for manuscript and art preparation are outlined on the inside back cover. Robert McC. Adams Secretary Smithsonian Institution SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY • NUMBER 60 Microdistribution of Foraminifera in a Single Bed of the Monterey Formation, Monterey County, California Roberta K. Smith and Martin A. Buzas SMITHSONIAN INSTITUTION PRESS City of Washington 1986 ABSTRACT Smith, Roberta K., and Martin A. Buzas. Microdistribution of Foraminifera in a Single Bed of the Monterey Formation, Monterey County, California. Smithsonian Contributions to Paleobiology, number 60, 33 pages, 4 figures, 2 plates, 7 tables, 1986.—While several papers exist on the small scale spatial distribution of living foraminifera, almost no work exists on the small scale spatial distribution of fossils. The present study took 24 (5 ml) replicates 10 cm apart along one bed of the Montery Formation in California. The mean density for all replicates is 6084.96 with a standard deviation of 8776.95. Both inspection and a cluster analysis of the data indicate replicates 20-24 have a much higher density and different rank order of abundance than replicates 1-19. The mean density for the total of all species in replicates 1-19 is 2387.47 with a standard deviation of 1175.58. For replicates 20-24 the mean density is 20135.40 with a standard deviation of 11181.40. The spatial variability is so great that four replicates (more than commonly taken) would only allow us to be 95% confident that we are within 50% of the true mean. Because age determination is based on presence of particular taxa rather than on densities, stratigraphic assignment would still be possible. The three species dominating the 1-19 group make up from 86% to 99% of the fauna. The three species dominating the 20-24 group make up from 77% to 85% of the fauna. Two of these are also dominant in the 1-19 group, but the most dominant species in the 20-24 group constitutes only <1% to 8% in the 1-19 group. The greatest number of species (22) occurs in the 20-24 group, as would be expected from the densities. The 1 — 19 group has 16 species. The infor¬ mation function is also highest in the 20-24 group. An attempt was made to achieve the faunal composition of the 1-19 group for replicates 20-24 by removal of percents of small-sized taxa. Comparable relative abundances are best achieved by removing 100% of Epistominella subperuviana and 95% of Bolivina brevior and other significant small-sized species. Total specimen numbers for both small- and large-sized species remains higher in replicates 20-24 than in 1-19, however. Thus, analysis of species percentages and species specimen size indicates that while transpor¬ tation—winnowing—of small specimens (or large specimens) into or out of the environment of deposition may be significant, it does not account for the differences between replicates 1-19 and 20-24. Therefore, either two habi¬ tats or some other mode of allocthonous enrichment or depletion rather than particle size winnowing must be invoked to account for the observed distri¬ bution. The low numbers of species, especially with so many individuals, indicates the fauna probably lived under stressful conditions. Low amounts of available oxygen may have caused the stress. Official publication date is handstamped in a limited number of initial copies and is recorded in the Institution’s annual report, Smithsonian Year. Series cover design: The trilobite Phacops rana Green. Library of Congress Cataloging in Publication Data Smith, Roberta K., 1931- Microdistribution of foraminifera in a single bed of the Monterey Formation, Monterey County, California (Smithsonian contributions to paleobiology ; no. 60) Bibliography: p. Supt. of Docs, no.: SI 1.30:60 1. Foraminifera, Fossil—California—Monterey County. 2. Paleontology—California Mon¬ terey County. I. Buzas, Martin A. II. Title. III. Series. QE701.S56 no. 60 560 s [563'. 12'0979476] 85-600357 [QE772] Contents Page Introduction. 1 Acknowledgments. 1 Methods. 2 Field. 2 Laboratory Sample Preparation Experiments. 2 Laboratory Preparation of 24 Replicates. 4 Microscope. 6 Results. 6 Replicates 1-24. 6 Density. 6 Replicates 1-19. 6 Replicates 20-24 . 7 Comparison of Replicates 1-19 and 20-24 . 7 Species Composition. 7 Replicates 1-19. 7 Replicates 20-24 . 11 Comparison of Replicates 1-19 and 20-24 . 12 Measurement of Species Diversity. 12 Species Diversity. 12 Replicates 1-19. 12 Replicates 20-24 . 13 Comparison of Replicates 1-19 and 20-24 . 13 Species Dominance Patterns and Species/Specimen Size. 14 Comparison of 24 Replicates with Subsamples from the Boulder Experiments. 14 Discussion. 15 Systematic Paleontology. 19 Literature Cited. 27 Plates. 29 m Microdistribution of Foraminifera in a Single Bed of the Monterey Formation, Monterey County, California Roberta K. Smith and Martin A. Buzas Introduction Several studies of small-scale spatial distribu¬ tion exist for living populations of benthic fora¬ minifera (e.g., Parker and Athearn, 1959; Buzas, 1965, 1968, 1970; Olsson and Ericksson, 1974). In general, these studies showed an inhomoge¬ neous distribution. A quantitative estimate of distributional variability is necessary before we can calculate confidence intervals for foramini- feral density or estimate the number of samples required for an arbitrarily chosen degree of con¬ fidence. The number of replicates required and the size or proximity of samples requires an understanding of small-scale spatial distribution. No direct way exists to pursue biology of fossil foraminifera; for paleobiology it is necessary to rely on work from living populations. Obviously, however, adequate sampling is also essential to paleoecological and paleoenvironmental recon¬ struction. Paleoecological work has lagged be¬ hind ecological in the area of sampling and small- scale spatial distribution. Roberta K. Smith, Earth Sciences Board, University of California, Santa Cruz, California 95064. Martin A. Buzas, Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Scott (1958) showed that fossil foraminifera were inhomogeneously distributed horizontally and vertically in an outcrop in New Zealand. At two stratigraphic levels in the Upper Tertiary of Maryland, foraminiferal species were homoge¬ neously distributed (Buzas and Gibson, unpub¬ lished). We know of no other small scale distri¬ bution studies of fossils. In the present study, we examined the small scale spatial distribution in a single bed—the same stratigraphic horizon—with evenly spaced replicates. We hoped to determine (1) the varia¬ bility among the replicates; (2) the confidence intervals for mean foraminiferal densities, and the confidence interval or precision obtainable for a given number of replicates; (3) the compa¬ rable adequacy of sampling for (a) time-strati¬ graphic, (b) broadly paleoenvironmental, and (c) paleoecological purposes; and (4) perhaps to draw some paleoecological conclusions about the sampled fauna and its environment. Acknowledgments. —Dr. Gu Gung-hsu and the Institute of Geophysics of the State Seismo- logical Bureau, Beijing, China, provided facilities for the extensive specimen count of the repli¬ cates. Micropaleontological laboratory facilities for sample preparation at the U.S. Geological 1 2 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY Survey, Menlo Park, California, were made avail¬ able to assistant Sandra Carlson by Drs. K. McDougal and R.V. Poore. Part of the technical assistance was funded through Dr. R.E. Garrison by the National Science Foundation (EAR 76- 2213). Drs. S.J. Culver, J.F. Evernden, and M. Gowing read and improved the manuscript, and Drs. J.F. Evernden and F.M. Govean gave other valuable assistance also. We are grateful to these persons and institutions. The term "clorox” is used throughout the text to indicate the commercial household bleach (so¬ dium hypochloride) that was used, full strength, during the laboratory work. Methods Field.— The Del Ray Canyon Diatomite member of the Monterey Formation is well ex¬ posed in a cut on Toro Road, near Monterey, California (see Figure 1). The bed studied by 24 replicate samples lies midway along the exposure, in unit 10 of Govean and Garrison (1981), ap¬ proximately 70 m (230 ft) stratigraphically above the base of the ash bed marking the base of the measured section. The boulder used in the pilot sample-preparation technique study came from 1 'A m stratigraphically above the 24-replicate bed. T he section shows a series of variously lami¬ nated and bedded to massive, softly diatoma- ceous to hard and cherty, mainly cream-colored marine sedimentary rocks. Externally massive- appearing but laminated, relatively thick beds of soft diatomaceous mudstone predominate. One of these was the source of the boulder used in the pilot study. The bed selected for the 24 replicate study is 8 cm thick and appears inter¬ nally finely shaly and laminated; it is grayish, weathering orange. It was chosen because (1) it is distinct from the more externally massive un¬ der- and overlying beds; (2) its thickness is ideal for the diameter of the coring device used; (3) it is soft enough to drive the coring device into; and (4) (a) it could be seen to contain foramini- fera, (b) it was believed that the fauna’s taxon¬ omic diversity would be relatively low, and (c) preservation appeared adequate to recover spec¬ imens from washed residue of the rock. The bed dips east for approximately 8 m diagonally across the road cut exposure from near the natural surface to the road bed level, approximately 3- 4 m elevationally lower. Twenty four replicate samples were taken 10 cm apart for 336 cm along the bed from the road bed level to a point a meter below the base of the soil profile (see Figure 1). The cut face inter¬ sects the bedding at 90 degrees, permitting hor¬ izontal penetration of the exposure with the cor¬ ing device. The coring device was a sharpened steel pipe with a 4 cm internal diameter welded to a steel rod and cross-bar. In spite of the diatomite’s softness and porosity, its considerable resistance only permitted driving the corer in 5 to 10 cm. Greater penetration could minimize possible surficial weathering effects. As the rock appears very porous, however, leaching may be general and not confined to surficial layers. After securing each replicate, it was extruded into a small sample jar. As the diatomite tended to fragment, we could not be sure to reject the possibly more weathered surface 2-3 cm. Laboratory Sample Preparation Experi¬ ments. —Various laboratories have experi¬ mented with simple to complex methods to ex¬ tract foraminifera from sediments and rocks, but the efficacy of these methods is not evaluated in the literature. We believe it is useful to include evaluation of preparatory methods because the very significant alterations of species densities which preparation techniques can affect can and do go unrecognized. These can just as seriously invalidate observations as can failure to sample adequately in the field. For this reason, sampling and preparation are treated together in this study. One reason we chose the soft diatomite was to capitalize on its ease of preparation. We made a pilot study to evaluate the effectiveness of the simplest laboratory preparation methods to ob- NUMBER 60 4 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY tain (nearly) all tests. The simplest techniques are to soak the material in various solvents, then rinse over sieves with water. Therefore, we subjected a number of subsam¬ ples from a diatomite boulder to soaking for two soaking periods—an arbitrarily chosen 6 months and 24 hours. No regard was paid to the boul¬ der’s stratigraphic setting or to spatial relations among subsamples. The first set of 10 subsamples and, later, a second set of 12 (plus one still later) were prepared from "slices” cut from large frag¬ ments of the boulder. The “slices” were broken to roughly 1 mm sizes (grains). For each subsam¬ ple increments of this debris were tamped down until the sediment in a 10-ml beaker reached 5 ml. Each of the first set of 10 5-ml volumes of sediment was soaked in 20 ml of 10 different solvents (with measured pH values) in closed 100- ml jars, at room temperature, for 6 months; another similar set was soaked for 24 hours. Solvents used were acetone, alcohol, carbon tet¬ rachloride, clorox, hydrogen peroxide, kero¬ sene, kerosene followed by water, mineral oil, distilled water, and unpurified tap water. For the 24-hour soak, a solution of 1 0% NaOH in distilled water was added. With both the 6- month and 24-hour soaks, microscopic sediment observations were made: immediately upon wet¬ ting; after 4 days; and after 6 months. Later, an additional subsample was prepared with “Qua¬ ternary O.” After soaking, each sediment subsample was rinsed over a sieve with 63 yum openings with warm tap water. Sieve residues washed onto filter papers were oven dried at 38°C. Subsequently, all of each residue was examined microscopically. The total specimen numbers for the 6-month and 24-hour soaks are shown in Tables 1 and 2. Most numbers from the 6 month soak are higher. An analysis of variance produced an Fj - = 0.55 value, which is, however, not significant. For the solvents tested (twice), short term soak¬ ing in kerosene then flushing with water is the most effective method. Long term clorox, hydro¬ gen peroxide, and mineral oil showed relatively good recovery also. The poorest specimen recov¬ eries are shown by carbon tetrachloride, water, plain kerosene, and sodium hydroxide solution (soaked only one day). Acetone shows an inter¬ mediate recovery. With all these solvents, many intact rock grains containing tests remained. As we wanted all tests freed, we next tried a more complex method. We followed manufacturer’s directions in a sev¬ eral-step procedure using the strong detergent “Quaternary O.” Recovery was significantly bet¬ ter but test-bearing rock fragments still re¬ mained. So we transferred our preparation work to the U.S. Geological Survey micropaleontology laboratory at Menlo Park and implemented their tested and more complex method for the 24 replicates, the data from which were used for the spatial study. Laboratory Preparation of 24 Repli¬ cates. —The method described below was adopted for all replicates in order to disaggregate (nearly) all of the diatomite to free all specimens, while minimizing mechanical and chemical test destruction. The method follows these steps. 1. Some fragmented diatomite was transferred from each field-sample jar to a 10-ml glass beaker. It was then tamped down, moistening slightly to assist compaction. This process was continued until the sediment was leveled at the 5-ml mark in the beaker. 2. Each 5 ml of sediment was placed in a 50- ml beaker and oven-dried overnight at 32°C. 3. The sediment in each beaker was covered with kerosene and left overnight. 4. The kerosene was decanted and the sedi¬ ment was covered to the 25-ml mark in the beaker with hot water; 2 ml of Na^CO^ (soda ash) were added. 5. Each sediment/liquid mixture was then boiled gently on an oscillating hot plate for ap¬ proximately one hour until break-down of the rock. 6. Material was rinsed through a 63 /im sieve with a small amount of detergent added to help remove the kerosene. Residues were collected on filter papers. 7. Residues in closed filter papers were oven NUMBER 60 bo c 15 cb o l/l s- 3 O _C I cm ~c c C3 C o u ,0 u QC LU 00 2 3 < OC LU LL < OC O O H co o co r^ lo cj> co co o co co 00 N. CO O) CM r-. o m o O) CM co o CO o O) CM CO CO co CM lO a; -O CO co a E o o _ O * • C0 co CD (/) 05 c CO 0) n E E o 0) OJ <0 a C/5 to CO o o +-> 0) c -Q 03 E O D » ® co ® C C a E £ k. O c ® s C (0 E £ CO 3 O to ® £ O CO C CO o At ® cn ® £ co £ E ® c <0 E c • c CO E £ 0) ® E <0 k. 03 c co E “5 •0 ** <0 £ TJ co E <0 CO c <0 o £ CO 3 O CO co c <0 > 3 CO o c k. o 3 k. co w a c ® 2 E • CO CO eo CO CO a c £ <0 3 o k. to «2 Q. (0 O >% 3 o <0 3 "S Q. c CO 3 O C 0 «M <0 m ■2 £ CO c co o> fll t 3 3 0 ) CO Q. o w o CO Q. d (0 P k, 03 a £ 3 to © o 3 i O d CO d CO ■ 4 C o c ® d CO c CO ® o CO c • E o o > a> k. £ <0 w 3 0 ) to a CO c c ’£ a> to «o I JC <0 ■0 c 3 k. 3 O <0 © c 3 ■o fO © c w 0) <0 © c 03 o CO c <0 c 1 3 £ co c k. ® c © 0 > co © C E o « c CO CO £ E ® £ c « C 3 ^ c «0 0> ^ 2 ® k. 5 o CO kk CO E o •c o to 2 ® c ? CO 0) c ® a CO o k. ® ® a CO O k. ® > > * O) o> | | 1 1 O £ o o £ o +* CO 1 ° 52 k* co o c o c £ E £ E o o o 3 3 3 3 3 t *>* d "5 <0 Q. o 0 3 □ 03 to 03 co 03 QQ Co 03 s <0 UJ 3 . UJ £ z z epunj66n$ * spuiuiA{|og spiufHJjni Sp|U|J3B|An v sp|U|ai||ng (l)BIJBUI01Std3 <8 spipiqjooeta sp|uo|uoN 6 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY Table 2.—Total numbers of tests recovered from 5 ml of sediment (ranked by 6-month abundance). Treatment 6 Months 24 Hours Soaked both periods Hydrogen peroxide 1037 574 Clorox 908 439 Kerosene-water 750 918 Acetone 525 411 Alcohol 344 308 Kerosene 290 308 Water-distilled 167 346 Carbon tetrachloride 152 116 Soaked only one period or otherwise not nu¬ merically comparable Water-industrial, old 318 166 (±'/3 spilled) Water-industrial, new - 2339 (different boulder fragment) Sodium hydroxide - 226 Mineral oil 729 - dried overnight at 32 °C. They were examined microscopically to determine if breakdown was adequate or if steps 2-7 needed repeating. Breakdown appeared adequate and they were retained for later investigation. Microscope.— All washed residues from all subsamples and replicates were examined under the microscope. All complete foraminiferal tests and fragments judged to represent individuals (mainly intact juvenile whorls) were picked from all subsamples and one replicate (no. 2) and mounted on slides. For all other replicates all such individuals were identified taxonomically and counted. Only specimens of taxa occurring rarely or requiring further study for identifica¬ tion were picked. Results Replicates 1-24.—Table 3 shows specimen counts for all 24 replicates. The mean for the total population is 6084.96 and the standard deviation 8776.95. The 95% confidence interval for the mean is given by A — 1.96(3%) represented. Taxa per subsample range from four to eight. One of the remaining three subsamples (24- hour soak in CC1 4 ) shows an unusual Nonionella NUMBER 60 15 dominance over Valvulineria (62% to 35%). An¬ other (6-month soak in mineral oil) represents a somewhat different fauna. In it taxa mainly are those above but dominance patterns differ, with greater species equitability and total number of taxa (10). Species percentages are (1) and (2) = 57%; plus (3) = 70%; (1) through (4) = 92%; also buliminaceans = 36%, (3) and (4) = 34%, and bolivinids = 7%. To replace one partly spilled subsample, an¬ other was prepared from a different boulder fragment. Although similar to the others, this fauna appears significantly denser and also tax- onomically resembles the fauna of replicates 20- 24. Taxa (1) and (2) = 41%; (3) = 22%; (4) is absent. Buliminaceans = 27%; 23% bolivines— four species, mainly Bolivina brevior Cushman; 5% Buliminella elegantissima d’Orbigny; and 2% Globobulimina pacifica Cushman. Neither the lat¬ ter two nor Epistominella subperuviana (Cush¬ man), here 7%, occur more than very rarely in other subsamples. Two other taxa (<1% each) complete the assemblage. In addition to the diatomite assemblages, two porcelanite thin sections from a few cm below the 24 replicate bed contain another perhaps distinguishable fauna (see Table 6), although specimen numbers are too low to be sure. We have, then, identified three to five distin¬ guishable faunas from this one exposure. We have done this in a study of 336 cm of one bed (the 24 replicates) and limited examinations of two or three other diatomite and porcelanite beds. We can compare these faunas in percents (Table 7), but differences in preparation meth¬ ods preclude comparison of densities. Other distinguishable faunas could be repre¬ sented as well. Some of the faunas described by Govean (1980) from the Toro Road stratigraphic section appear distinct. Time stratigraphic signif¬ icance may be nil, but paleoecological signifi¬ cance may be considerable. Discussion The 24 replicate samples enabled us to docu¬ ment the variability of fossil foraminifera in the horizon studied. Studies of spatial distribution of modern foraminifera (Buzas, 1968, 1970;Olsson and Eriksson, 1974) indicate living and dead populations are inhomogeneously distributed. The very large and abrupt change observed herein, however, has never before been recorded in either a modern or a fossil population. Unfor¬ tunately, very few studies documenting micro¬ distributions exist. Whether or not we are ob¬ serving a bizarre phenomenon in the present study cannot be ascertained until more studies are made. Normal paleontological sampling would not detect the changes observed herein. The very high densities observed in this study are seldom recorded in living populations. Dens¬ ities as high as about 4000 per 5 ml were re¬ corded in caging experiments (Buzas, 1978), and Sen Gupta et al. (1981) recorded living densites of 3132 in 3 ml of sediment on the continental slope off Daytona Beach, Florida. Interestingly, the most abundant species recorded by Sen Gupta et al. (1981) belonged to the genus Boli¬ vina. Usually, densities are in the tens or hundreds per 5 ml for living populations. Even for total populations, densities of thousands and tens of thousands are seldom recorded. They do, however, occur (see for example, Phleger, 1951; Buzas, 1965). We are, then, probably observing in this fossil population the accumulation of tests over some period of time. Traditional, though undocumented, micropa- leontological sampling techniques may have taken into account specimen patchiness—both horizontal and vertical. These techniques include hand lens examination of rocks in the field. Such examination reveals some concentrations of for¬ aminifera on bedding planes, scattered to con¬ centrated foraminifera in areas and volumes of rocks, and sparsely populated or barren rock. The intent of the examination is to assure that fossiliferous rocks are collected—not to docu¬ ment distribution. The assumption has been that whatever is collected will be “representative”— especially when rock ages primarily are sought. While this assumption is probably true relative to age, it is far less true relative to paleoecology. Distribution and abundance provides the frame- 16 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY Table 5.—Replicates 20—24, species numbers and percentages adjusted in attempt to achieve the faunal composition of replicates 1-19. Various totals Taxa reduced by 0% Replicate number Total in replicate Total all except Valvulineria, B. curta, and Nonionella Total Valvulineria cf V. c. obesa, B. curta, and N. schencki Total “ Bolivina ’’ including Suggrunda Other taxa Other buliminaceans CAobobulimina pacifica Bolivina seminuda Nonionella schencki 12448 8467 3981 7710 10 10 27 142 492 100% 68% 32% 62% <1% <1% <1% 1% 4% 6156 2175 (3981) 2034 (10) (10) (27) (142) (492) O A (100%) 35% 65% 33% <1% <1% <1% 2% 8% zu 4965 984 (3981) 899 (10) (10) (27) (142) (492) (100%) 20% 80% 18% <1% <1% 1% 3% 10% 4568 587 (3981) 520 (10) (10) (27) (142) (492) (100%) 13% 87% 11% <1% <1% 1% 3% 11% 39561 19806 19755 15188 7 87 126 552 2193 100% 50% 50% 39% <1% <1% <1% 1% 6% 24674 4919 (19755) 4211 (7) (87) (126) (552) (2193) 9 1 (100%) 20% 80% 17% <1% <1% 1% 2% 9% Z 1 22180 2425 (19755) 2016 (7) (87) (126) (552) (2193) (100%) 11% 89% 9% <1% <1% 1% 2% 10% 21357 1602 (19755) 1284 (7) (87) (126) (552) (2193) (100%) 7.5% 92.5% 6% <1% <1% 1% 3% 10% 16783 8583 8200 6871 5 10 10 351 965 100% 51% 49% 40% <1% <1% <1% 2% 6% 10412 2212 (8200) 1981 (5) (10) (10) (351) (965) 99 (100%) 21% 79% 19% <1% <1% <1% 3% 9% ZZ 9310 1110 (8200) 1003 (5) (10) (10) (351) (965) (100%) 12% 88% 11% <1% <1% <1% 4% 10% 8944 744 (8200) 677 (5) (10) (10) (351) (965) (100%) 8% 92% 8% <1% <1% <1% 4% 11% 13006 7722 5284 6657 0 5 14 136 699 100% 59% 41% 51% 0% <1% <1% 1% 5% 7176 1892 (5284) 1766 (0) (5) (14) (136) (699) o<2 (100%) 26% 74% 25% 0% <1% <1% 2% 10% 6134 850 (5284) 788 (0) (5) (14) (136) (699) (100%) 14% 86% 13% 0% <1% <1% 2% 11% 5187 503 (5284) 462 (0) (5) (14) (136) (699) (100%) 9% 91% 8% 0% <1% <1% 2% 12% 18879 12305 6574 10001 5 35 31 229 712 100% 65% 35% 53% <1% <1% <1% 1% 4% 941 1 2837 (6574) 2435 (5) (35) (31) (229) (712) (100%) 30% 70% 26% <1% <1% <1% 2% 8% Z4 7984 1410 (6574) 1206 (5) (35) (31) (229) (712) (100%) 18% 82% 15% <1% <1% <1% 3% 9% 7429 855 (6574) 718 (5) (35) (31) (229) (712) (100%) 11.5% 88.5% 10% <1% <1% <1% 3% 10% NUMBER 60 17 Table 5.—Continued. Taxon reduced by 100% Taxa reduced by 75%, 90%, and 95% Buliminella curta Valvulineria cf. V. californica obesa Epistominella subperuviana Oolina, Lagena, and Nodosaria? Buliminella elegantissima Suggrunda kleinpelli Bolivina brevior ar other small Bolivia 1517 1972 396 5 370 423 7145 Original No. — 100% 12% 16% 3% <1% 3% 3% 57% % of Fauna (1517) (1972) 0 1 93 106 1786 No. Reduced 75% 25% 32% 0% <1% 2% 2% 29% % of Fauna (1517) (1972) 0 1 37 42 715 No. Reduced 90% 31% 40% 0% <1% 1% 1% 14% % of Fauna (1517) (1972) 0 <1 19 21 357 No. Reduced 95% 33% 43% 0% <1% <1% <1% 8% % of Fauna 7003 10559 2170 78 1871 1676 12960 Original No. — 100% 18% 27% 5% <1% 5% 4% 33% % of Fauna (7003) (10559) 0 20 468 419 3240 No. Reduced 75% 28% 43% 0% <1% 2% 2% 13% % of Fauna (7003) (10559) 0 8 187 168 1296 No. Reduced 90% 32% 48% 0% <1% 1% 1% 6% % of Fauna (7003) (10559) 0 4 94 84 648 No. Reduced 95% 33% 49% 0% <1% <1% <1% 3% % of Fauna 2781 4454 743 10 810 600 5920 Original No. — 100% 17% 27% 4% <1% 5% 4% 35% % of Fauna (2781) (4454) 0 3 203 150 1480 No. Reduced 75% 27% 43% 0% <1% 2% 1% 14% % of Fauna (2781) (4454) 0 1 81 60 592 No. Reduced 90% 30% 48% 0% <1% 1% 1% 6% % of Fauna (2781) (4454) 0 1 41 30 296 No. Reduced 95% 31% 50% 0% <1% <1% <1% 3% % of Fauna 1803 2782 620 11 416 324 6197 Original No. — 100% 14% 21% 4% <1% 3% 2% 48% % of Fauna (1803) (2782) 0 3 104 81 1549 No. Reduced 75% 25% 39% 0% <1% 1% 1% 22% % of Fauna (1803) (2782) 0 1 42 32 620 No. Reduced 90% 29% 45% 0% <1% 1% 1% 10% % of Fauna (1803) (2782) 0 1 21 16 310 No. Reduced 95% 31% 48% 0% <1% <1% <1% 5% % of Fauna 2199 3663 908 16 1307 1058 8714 Original No. — 100% 12% 19% 5% <1% 7% 6% 46% % of Fauna (2199) (3663) 0 4 327 265 2179 No. Reduced 75% 23% 39% 0% <1% 3% 3% 23% % of Fauna (2199) (3663) 0 2 131 106 871 No. Reduced 90% 28% 46% 0% <1% 2% 1% 11% % of Fauna (2199) (3663) 0 1 65 53 436 No. Reduced 95% 28% 49% 0% <1% 1% 1% 5% % of Fauna 18 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY Table 6. —Faunal composition and rank for two thin sections from porcelanite bed. Abundance Rank of species Abundant 1. Valvulineria cf. V. californica obesa Common-abundant 2. Bolivina brevior and other small bolivinids, including Suggrunda 3. Buliminella curta Few-common 4. Bolivina seminuda Few 5. Globobulimina pacifica 6. Nonionella schencki Rare-few Rare 7. Buliminella elegantissima 8. Epistominella subperuviana 9. Bulimina cf. B. pseudoaffinis (small) work for paleoecology as it does for ecology. We have herein established that one fauna could not be derived from the other by simple transportation. How closely the fossil population resembles the living populations of the Miocene sea floor is difficult to evaluate. We cannot be sure if we are witnessing a true change in the fauna due to some abiotic or biotic change or if the fauna was transported from somewhere else or both. Any and all replicates provide for the same assignment of Mohnian Age (of Kleinpell, 1938) based on the presence of Nonionella schencki. Similarly, any and all provide for the same pa- leoenvironmental interpretation of “medium depths,” “probably upper bathyal.” Govean (1980) and Govean and Garrison (1981) have described the Toro Road section as forming near the top of the bathyal zone at 150 to 500 m. Note that this classical sort of paleoenvironmen- tal interpretation concerns itself primarily with broad-scale depth ranges (relating to basin recon¬ struction). Such other (actual paleoecological) variables as temperature, salinity, and available oxygen, as well as redeposition, are also consid¬ ered where evidence appears to exist for their interpretation. In this case, it is reasonable to assume “cool temperatures” and “normal marine salinity.” This is so even though the species diversity is fairly low, a condition suggesting some sort of stress situation for foraminifera as a group. No foraminifera thought to represent either high or low or variable salinity ranges or particularly warm or cold temperatures were found. Regarding available oxygen, Govean (1980) and Govean and Garrison (1981) have inter¬ preted some parts of the stratigraphic sequence exposed on Toro Road as representing “oxygen- minimum” conditions. The present replicates showing a relatively low species diversity would lend themselves to that (stress) interpretation. They do not, however, contain abundant speci¬ mens of taxa specifically interpreted as repre¬ senting 0 2 minima, although Bolivina seminuda and Suggrunda occur (see Phleger and Soutar, 1973; Byers, 1977; Ingle et ah, 1980). Overall, the faunal composition would not necessarily be taken to indicate 0 2 minima, although it may indicate somewhat reduced 0 2 conditions. On the other hand, the fine laminations seen particularly well developed within the bed sam¬ pled are also believed to indicate 0 2 minima. This is because lamina develop and remain undis¬ turbed where low oxygen concentrations pro¬ hibit burrowing organisms that would disrupt laminae. The Valvulineria specimen size (small for the V. californica group) may reflect 0 2 minima or, simply, stress conditions for many foraminifera; a “population explosion’’ of small specimens of a taxon may result from the absence of competi¬ tion or predators. Another explanation is that the small area represented by replicates 20-24 was for some reason extremely good for forami¬ nifera. On the Mississippi Delta, Lankford (1959) found where foraminifera were most abundant the tests were the smallest. We also observed that the smallest tests occur where the densities are the highest. Perhaps there was a population ex¬ plosion that correlated with variation in 0 2 con¬ tent. NUMBER 60 19 Systematic Paleontology Herein the classification of Loeblich and Tap- pan (1964, 1974) is followed, with some modifi¬ cation. All commoner taxa have been compared with types erected by Kleinpell (1938) and de¬ posited in the micropaleontology museum collec¬ tions of Stanford University. Access to these types was kindly provided by Dr. J.C. Ingle. Some preliminary identifications were made by Dr. F.M. Govean of AMOCO Production Company, Tulsa, Oklahoma. Unfortunately, most speci¬ mens of rarely occurring taxa were lost in transit between Tulsa and Santa Cruz, California, pre¬ venting their comparison with types. Order Foraminiferida Eichwald, 1830 Family Nodosariidae Ehrenberg, 1838 Genus Nodosaria Lamarck, 1812 Genus Lagena Walker and Jacob, 1798 Family Glandulinidae Reuss, 1860 Genus Oolina d’Orbigny, 1839 Family Turrilinidae Cushman, 1911 Genus Buliminella Cushman, 1911 Family Bolivinitidae Cushman, 1927 Genus Bolivina d’Orbigny, 1839 Genus Suggrunda Hoffmeister and Berry, 1937 Family Buliminidae Jones, 1875 Genus Bulimina d’Orbigny, 1826 Genus Globobulimina Cushman, 1927 Family Uvigerinidae Haeckel, 1894 Genus Siphogenerina Schlumberger, 1883 Genus Trifarina Cushman, 1923 Genus Uvigerina d’Orbigny, 1826 Family Discorbidae Ehrenberg, 1838 Genus Epistominella Husezima and Maruhasi, 1944 Genus Valvulineria Cushman, 1926 Family Epistomariidae Hofker, 1954 Genus Epistomaria Galloway, 1933 Family Nonionidae Schultze, 1854 Genus Nonionella Cushman, 1926 Family Anomalinidae Cushman, 1927 Genus Holmanella Loeblich and Tappan, 1962 Nodosaria? sp. Four broken nodosarine specimens, each with two elongate chambers, were found. They closely resemble Nodosaria parexilis Cushman and Stew¬ art (in Stewart and Stewart, 1930) or N. tympan- iplectriformis Schwager of Haller (1980:235, pi. 3: fig. 10), identified from Pliocene beds near the northern California coast. Hypotype: USNM 382514. Lagena sp. Three inornate specimens belong to Lagena. ?Oolina globosa (Montagu) \^']Oolina globosa (Montagu).—Kleinpell, 1938:225 (= La¬ gena globosa Montagu). This inornate Oolina is consistently present in replicates 20-24, totaling approximately 125 specimens. Hypotype: USNM 382515. Buliminella curta Cushman Plate 1: figures 1, 2 Buliminella curta Cushman, 1925:33, pi. 5: fig. 13; 1926:55.—Kleinpell, 1938:248, pi. 7: fig. 4, pi. 16: fig. 8 . Thirty Stanford University collections hypo- types from the Salinas shale and Modelo, Tem¬ blor, Monterey, and possibly other formations have been examined. Most of the abundant pres¬ ent specimens had the final few large chambers fragmented or broken off from earlier whorls (probably in sample preparation). This makes the population superficially appear to be composed of relatively small specimens. Yet, it still is clear that the population shows considerable range of variation in specimen height/breadth—as do the Stanford hypotypes. Figured Hypotypes: USNM 387632, 387633. Hypotype: USNM 382516. Buliminella dubia Barbat and Johnson Buliminella dubia Barbat and Johnson, 1934:13, pi. 1: figs. 14, 15.—Kleinpell, 1938:249, pi. 16: fig.7. Two small Toro Road specimens (one each from alcohol and clorox six-month soaks) com- 20 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY Table 7.—Check list for all observations. ■ abundant (33-100%) 24 REPLICATE SAMPLES FROM ONE BED • COMMON (12-33%) O FEW (3-12%) A RARE (< 1-3%) Butiminaceans Boiivinitids & Suggrunda Bolivina brevior Bolivina seminuda Bolivina spp. (4) Suggrunda kleinpelli Turrilinids Buliminefla curt a Buliminella dubia Buliminella elegantissima Buliminids Bulimina cl. B. pseudoaffinis Globobulimina pacifica Uvigerinids Siphogenerina sp. Trifarina sp(p). Uvigerina sp(p). Cassidulinaceans Nonionids Nonionella schencki Discorbaceans Discorbids Epistominella subperuviana Valvulineria cf. V. californica obesa Valvulineria (?) sp. cf. V. araucana Nodosariaceans Glandulinids Oollna sp. Lagenids Lagena sp. A Nodosaria (?) sp. Others (5 spp.) TOTAL NO. SPECIES IN REPLICATE /SUBSAMPLE TOTAL NO. SPECIMENS IN REPLICATE /SUBSAMPLE 1 2 3 4 5 6 7 8 9 10 1 1 12 13 14 15 16 17 18 19 20 21 22 23 24 o A A A o A Cs A A o o O o o o o A A o ■ ■ ■ ■ ■ o A A A A A o O o o A o 0 A A A Cl A o A A A A A A A A A A A A A Cl A A A A A A A A A A A A A A A A A A A A Cl A A A o O o A O • A • A • A • • • Cl • A • • • A • • A ■ ■ a • a • A • • A • A • A • A • A • A • A A A Cl A A A A A cs Cl A A A A A A o o o O o A A o A A A A A o A A o A A A A A A A A Cl A Cl Cl A A A A A A A A A A #••■•■■•■•••••■••00000 00000 A AAaAaaaA A A A A A A A A A A 1 1 9 10 10 10 8 10 9 8 9 7 1 1 12 12 13 13 6 1 1 14 19 20 16 16 19 CO CM CD COCOCOOOJOOO^Cg CO O 00 ID CO CO CO ® »-o co r*. Or>-.o© ^ CO CM CM CM © CM ® (D CO © r- CO r- r- pared favorably with the holotype and paratypes and the specimen figured by Kleinpell, all from Upper Miocene rocks, in the Stanford collec¬ tions. From the specimens seen, this form is nearly bulimine in coiling—barely if exceeding three chambers per whorl. Hypotype: USNM 382517. Buliminella elegantissima (d’Orbigny) Plate 1: figure 3 Bulimina elegantissima d’Orbigny, 1839:51, pi.7: figs. 13, 14. Buliminella elegantissima (d’Orbigny).—Kleinpell, 1938:249, pi. 16: fig. 10.—Smith, 1978:141, pi. 2: fig. 1.—Buzas, Smith, and Beem, 1977:71, pi. 1: figs. 19, 20. Twelve Miocene hypotypes in the collections of Stanford University ascribed to the species by Kleinpell (1938) and many hypotypes at the Na¬ tional Museum of Natural History (Smith, 1978; Buzas, Smith, and Beem, 1977; and others) have been examined. Like d’Orbigny’s, most USNM types are Recent, but the species seems little changed since the Paleocene. Perhaps detailed time-morphology studies would reveal evolution¬ ary patterns. Figured Hypotype: USNM 387634. Hypotype: USNM 382518. NUMBER 60 21 Table 7.—Continued. SUBSAMPLES FROM ONE BOULDER SOAKED IN DIFFERENT SOLUTIONS 6 MONTH SOAKING TIME 24 HOUR SOAKING TIME *6 O “ •a CM T? c X © © c c CO a © o X © © Q co Z c. o CM 0) 0J X o O O o o O o o o O o CM © © ffl CM CM CM < O O X * * z X I I PORCELANITE THIN SECTIONS 2 A A A A A A A o A A o A A A A A o o A O o o o • o O O o A A A A A o o A o o o o • o o o o A A A A • • A A O O A A A A A A A OOOOAAO* ■■ ' A o o o o o o o o o o AAA A A o A A O A A O O A 8 8 co o 6 10 CM r- A 5 12 co o co co ^ CD ffl r*. o i- co O) <0 CO CD CM 4^ CO CM <0 r- a> CO co CM A Bolivina brevior Cushman Plate 1: figures 4-8 Bolivina brevior Cushman, 1925:31, pi. 5: fig. 8a,b; 1926:54.—Kleinpell, ed., 1980, pi. 7: fig. 11. Bolivina brevior brevior Cushman.—Kleinpell and Tipton, 1980:72. species occurs in all the material studied and dominates replicates 20-24 with thousands of specimens. Figured Hypotypes: USNM 387635- 387637. Hypotype: USNM 382519. California hypotypes examined in the Stanford collections are (1) three “Upper Miocene’ from a Kettleman Hills well; (2) one “Miocene” from the Salinas shale, Reliz Canyon, Monterey County; (3) two from the Gould shale, Zemorra Creek, Kern County; (4) and four from the Mon¬ terey formation, Santa Barbara County. This ?Bolivina conica Cushman [}]Bolivina conica Cushman, 1925:30, pi. 15: fig. 4a,b; 1926:54.—Kleinpell, 1938:269, pi. 7: fig. 7a,b. A single, large, conical, costate specimen is so referred. 22 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY Bolivina dunlapi Kleinpell Bolivina dunlapi Kleinpell, 1938:271, pi. 15: fig. 2. Bolivina brevior dunlapi Kleinpell.—Kleinpell and Tipton, 1980:72. The holotype was examined. Kleinpell and Tipton (1980) state that “except for its costae, this small form is very similar to Bolivina brevior. Bolivina dunlapi Kleinpell is herein reinterpreted as the costate subspecies of B. brevior. 7 ' We, how¬ ever, presently retain B. dunlapi as a separate species to which a few specimens from Toro Road seem best referred, although they also re¬ semble B. sulphurensis Cushman and Adams. Hypotype: USNM 382520. Bolivina pseudospissa Kleinpell Plate 1: figure 13 Bolivina pseudospissa Kleinpell, 1938:279, pi. 21: fig. 6.— Kleinpell, ed., 1980, pi. 7: fig. 4. A few, distinctive, compressed specimens with very neatly arranged nonlobate arcuate sutures seem best referred to this species. Compared with the holotype, they have the same chamber and suture pattern and considerable degree of com¬ pression but lack a keel. Populations of Bolivina pseudospissa were not available for comparison, however. These Toro Road Bolivina also closely resemble B. paula Cushman and Cahill from the Pliocene Yorktown formation of the East Coast. Figured Hypotype: USNM 387641. Hypotype: USNM 382521. Bolivina rankini Kleinpell Bolivina rankini Kleinpell, 1938:288, pi. 22: figs. 4, 9. A few specimens from Toro Road appear very similar to the holotype, but distinct from Bolivina seminuda Cushman, being more tapering and compressed. They appear best referred to this species. Hypotype: USNM 382522. Bolivina seminuda Cushman Plate i: figures 9-12 Bolivina seminuda Cushman, 1911:34, fig. 55. Kleinpell, 1938:281. Bolivina seminuda Cushman forma seminuda Govean, 1980:146, pi. 1: figs. 1-5, pi. 3: figs. 2-6, pi. 4: figs. 1-6, pi. 5: figs. 3, 4, 6, pi 6: figs. 4, 5, pi. 7: figs. 1-6, pi. 8: figs. 1-5, 7, pi. 9: figs. 1-3, pi. 10: figs. 1-5. Bolivina seminuda seminuda Cushman.—Kleinpell, ed., 1980, pi. 8: figs. 5, 6, 9, 10. Bolivina foraminata R.E. Stewart and K.C. Stewart. Klein¬ pell, ed., 1980, pi. 8: figs. 7, 8. Bolivina seminuda Cushman subspecies foraminata Stewart and Stewart.—Govean, 1980:145, pi. 2: figs. 1-3, pi. 3: fig. 1, pi. 5: figs. 1, 2, 5, pi. 6: figs. 1-3, pi. 7: fig. 7, pi. 8: fig. 6. The 16 Stanford hypotypes referred to Boli¬ vina seminuda and the 12 referred to B. seminuda foraminata by Kleinpell (1938:281) were exam¬ ined. Govean (1980) showed that the B. seminuda and B. foraminata forms are ecophenotypes. The species was found throughout the Toro Road material studied but in relatively small numbers. Figured Hypotypes: USNM 387638- 387640. Hypotype: USNM 382523. Suggrunda kleinpelli Bramlette Plate 1: figures 14-17 Suggrunda kleinpelli Bramlette, in Woodring and Bramlette, 1950:59, pi. 23: figs. 4, 5, 9. The present specimens, numbering several hundred, clearly belong to Suggrunda. Many closely resemble Bramlette’s type Figures. The holotype comes from a “road cut on Laureles grade,” very near the present location and ap¬ parently from the same stratigraphic unit. Yet, much variation occurs in the present population of Suggrunda —as to (1) quadrateness, (2) com¬ pression, (3) marginal sharpness, and (4) spine development—four related characters. Deter¬ mining whether or not distinct morphological groups and possibly taxa are represented would require further study. Populations of Bramlette’s NUMBER 60 23 form also should be examined. Suggrunda was placed in the Caucasinidae of the Cassidulinacea by Loeblich and Tappan (1964), but is retained in the Bolivinitidae herein. Figured Hypotypes: USNM 387642, 387643. Hypotype: USNM 382524. Bulimina cf. B. pseudoaffinis Kleinpell Plate 2: figures 1, 2 This form commonly constitues from 1 %- 10% of the assemblages from the boulder subsamples, but was not found in any of the 24 replicates from one bed. Kleinpell’s holotype was exam¬ ined, but no other specimens were seen in the Stanford collections. The holotype is preserved differently than the present specimens—giving a different appearance. Its apertural area also ap¬ pears to have been somewhat squashed. This may give this specimen the “thickest near middle” outline described by Kleinpell (1938:257, pi. 9: fig. 9). If that characteristic represents his popu¬ lations, however, it may not match the present specimens; they are thickest from middle to up¬ per third. Their sutures also appear a bit more depressed than Kleinpell’s holotype, but in. the absence of a population, it is not possible to know certainly if this is true. Our specimens also appear a bit smaller than the holotype of B. pseudoaffinis ; this could be environmental, however. Kleinpell had originally identified the holotype as a member of B. affinis d’Orbigny and re¬ marked (1938:258) that B. pseudoaffinis is "ap¬ parently closely related to’ that taxon. Interest¬ ingly, the figures given by Haller (1980:246, pi. 7: fig. 6a,b) for Globobulimina affinis (d’Orbigny) (from the Pliocene Rio Del Formation) very closely resemble the present specimens. No Haller specimens were seen, but perhaps the Toro Road form is intermediate and B. pseu¬ doaffinis is the ancestor. As to generic identity, the present specimens could be placed in either Bulimina or Praeglobulimina or Globobulimina on the basis of the degree of overlap/envelopment of chambers. The condition of the specimens studied herein did not allow for investigation of tooth-plate characteristics. Figured Hypotypes: USNM 387644, 387645. Hypotype: USNM 382525. Globobulimina pacifica Cushman Plate 2: figure 3 Globobulimina pacifica Cushman, 1927:67, pi. 14: fig. 12a,b.—Kleinpell, 1938:260, pi. 8: fig. 7. Both the holotype and Kleinpell’s figured spec¬ imen have been examined. This species is well represented throughout the Toro Road mate¬ rials, although never common. Most tests were broken in sample preparation. Numerical abun¬ dances were estimated from fragmentary speci¬ mens. Figured Hypotype: USNM 387646. Hypotype: USNM 382526. Siphogenerina sp. A single early-test portion is questionably re¬ ferred to this genus. It was found in the distilled water one-day-soak boulder subsample. It is ro¬ bust, nearly rounded, and has sutural lobation; it is biserial. It could be a Bolivina but seems better referred to Siphogenerina. Trifarina sp(p). Trifarina is represented by two costate and five smooth specimens in this Toro Road material. Other than ornamentation, they are very similar. Such ornamented and unornamented Trifarina may belong to more than one species. Uvigerina spp. Uvigerina is represented by fewer than 20 spec¬ imens in the Toro Road material. These all are 24 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY large; most are smooth and one generically ques¬ tionable specimen is costate. Hypotype USNM 382527 represents the smooth variety. Hypotype: USNM 382527. Epistominella subperuviana (Cushman) Plate 2: figures 4-6 Pulvinulinella subperuviana Cushman, 1926:63, pi. 9: fig. 9. The holotype is deposited in the National Mu¬ seum of Natural History. A specimen in the Stanford collection (LSJU type no. 943, slide 1045) might be a paratype (see Cushman, 1926:63) but was figured by Kleinpell (1938:321, pi. 14: fig. lOa-c) as Eponides sp. It is from the Salinas shale, Monterey County, California. This specimen and two hypotypes also identified by Kleinpell as Eponides sp. have been examined and appear to be Epistominella subperuviana. These two hypotypes are from the upper type Monterey Formation and the Monterey Forma¬ tion of the Nipomo Quadrangle, respectively. A very similar species is Epistominella relizensis (Kleinpell). The holotype of Pulvinulinella reli¬ zensis Kleinpell (1938:329, pi. 10: fig. lOa-c) is missing. Only one other specimen so ascribed is in the Stanford colections. Yet, (1) the type de¬ scriptions of “P. ” relizensis and “P. ” subperuviana and (2) comparison with the one specimen as¬ cribed to the former and the three ascribed to Eponides sp. by Kleinpell indicate that the Toro Road specimens are probably not best referred to Epistominella relizensis but to E. subperuviana. Epistominella relizensis is described with a smaller holotype than E. subperuviana. The smaller size is in keeping with the present specimens, but otherwise they are more like E. subperuviana. (Note that some Toro Road species—as Valvuli- neria —are smaller than elsewhere.) Kleinpell (1938) stated that the “test of P[ulvinulinella] relizensis is more strongly and symmetrically biconvex than in P subperuviana, typical specimens of which also are present in Reliz Canyon.” The present specimens have the test approximately as biconvex as does Epistomi¬ nella subperuviana —not more. The one type of E. relizensis in the Stanford collections does show a more biconvex form. We compared Kleinpell’s (1938:329, pi. 16: fig. 5a—c) specimen of ” Pulvinulinella ” relizensis to another from Reliz Canyon (Lower Delmon- tean part of the section) ascribed to and figured as P. cf. P pontoni Cushman. They are very similar except that the “P.” relizensis specimen has flush sutures. Populations can, however, show both flush and depressed sutures. Five specimens of the form referred to P cf. P. bradyana Cushman by Kleinpell (1938:327) were examined also. The sutures range from flush to slightly depressed. They also closely re¬ semble the “P ” relizensis specimen. They are from the “Upper Modelo” Formation in Los Angeles County—“Lower Delmontean.” Thus, this form and “P.” cf. P. pontoni (above) are younger than the Kleinpell specimens of “P.” subperuviana and “P ” relizensis (Relizian and Lui- sian). The Toro Road specimens are of Mohnian age. Figured Hypotypes: USNM 387647, 387648. Hypotype: USNM 382528. Valvulineria sp. cf. V. araucana (d’Orbigny) A few specimens, distinct from Valvulineria cf. V. californica subsp. obesa, are so referred. They are similar to “ V. araucana (d’Orbigny) var. mal- agaensis" Kleinpell (1938:308, pi. 22: figs. 10- 12 ). Hypotype: USNM 382529. Valvulineria cf. V. californica Cushman subsp. obesa Cushman Plate 2: figures 7-12 Valvulinerias in the Stanford University type collections include many specimens pertinent to the problem of identifying the abundant Toro Road form. The Stanford types include para- types and hypotypes (and “plesiotypes”) ascribed by Kleinpell (see Kleinpell, 1938) and others to the taxa discussed below. NUMBER 60 25 Valvulineria miocenica Cushman (1926:61, pi. 8: figs. 9,10, pi. 9: fig. 3a-c; Kleinpell, 1938:313, pi. 61: fig. la-c) clearly is more compressed than the Toro Road Valvulineria and has sutures much more curved and nearly flush and thus has a less lobate periphery. Like V. miocenica, V. californica californica Cushman (V. californica Cushman, 1926:60, pi. 9: fig. la-c; Kleinpell, 1938:308, pi. 13: fig. 6a- c, pi 16: fig. 4a-c) and V. grandis Cushman and Galliher (1934:26, pi. 4: fig. 12a-c; Kleinpell, 1938:312) are more compressed than the Toro Road form. Valvulineria californica appressa Cushman (V. californica Cushman var. appressa Cushman, 1926:60, pi. 9: fig. 5a-c; Kleinpell, 1938:309, pi. 13: fig. 7a—c) and V. californica obesa Cushman (V. californica Cushman var. obesa Cushman, 1926:61, pi. 9: fig. 2a-c; Kleinpell, 1938:310, pi. 10: fig. 12a-c, pi 14: fig. 12a-c) are not more compressed. Yet they are larger and appear to have less depressed sutures and consequently much less lobate periphery than the Toro Road form. Marginal sutural depression is so great with the later chambers of the Toro Road specimens than many nearly appear to approach “uncoil¬ ing.” They also have thinner walls than these V. californica types from Stanford. They do not, however, appear to closely resemble any other Valvulineria species. The Toro Road Valvulineria come from a very well studied formational and time-stratigraphic unit. It seems unlikely that they represent a pre¬ viously undescribed species or even subspecies. Although, if subspecies is conceived as applicable or relative to a geographic or ecologic morpho¬ logical (evolutionary) adaptation, a "new” subspe¬ cies could/may be represented. The most likely explanation of the morphol¬ ogy of the Toro Road Valvulineria follows. (1) The thinness of the wall—relative to character¬ istic and typical V. californica —reflects lesser availablility of Ca ++ + C0 3 in the life environ¬ ment. (2) The “thinness” also expresses itself as less sutural filling, giving more depressed (ap¬ pearing) sutures and lobate margin. (3) This “thinness’ and apparent marginal sutural depres¬ sion “culminate” in the final chambers appearing almost detached from the earlier whorl. It is possible, however, that an evolutionary trend toward extreme sutural depression—and even separation— is represented herein. (4) Relatively small specimen size may also reflect the life en¬ vironment. One explanation is that an abun¬ dance of small specimens represents rapid prolif¬ eration of a taxon in an optimum environment, or one without predators or competitors. All of these (kinds of) morphological factors have been correlated with some environmental stress conditions for foraminifera. In the present case, these could correlate with an oxygen-mini- mum environment (Ingle, pers. comm., 1983). Whether or not a new subspecies is represented here seems moot, but at present it also seems best to refer these specimens tentatively to Val- vulinera californica subsp. obesa. A study of the distribution of the Toro Road form is needed before this taxonomic problem can be resolved. Figured Hypotypes: USNM 387649- 387654. Hypotype: USNM 382530. Epistomaria sp. One specimen from a boulder subsample is so referred. Nonionella schencki (Kleinpell) Plate 2: figures 13-16 Nonion schencki Kleinpell, 1938:235, pi. 16: fig. lla,b.— Kleinpell, ed., 1980, pi. 2: fig. 2, pi. 3: figs. la,b, 2a,b, 5. The Toro Road specimens appear conspecific with the holotype from “4 mi. E. of Del Monte, Monterey County, California.” The paratype from the “Salinas Shale—Miocene” is larger than most of the abundant present specimens but has a very good likeness. Two hypotypes identified by Kleinpell from the “Santa Margarita Shale . . . Nipomo Quadrangle” are similar to the Toro Road form but preservation differs. Nine of Kleinpell’s (1938) specimens from the “Upper type Monterey” also show a very good likeness 26 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY to the Toro Road form. These types are in the Stanford collections. It is not entirely clear whether this form should be placed in Nonionella or Florilus. Figured Hypotypes: USNM 387655- 387658. Hypotypes: USNM 382531-382533. Nonionella sp. Two specimens are so referred; they are not Nonionella schencki (Kleinpell). Hypotype: USNM 382534. Holmanella sp. cf. H. valmonteenis (Kleinpell) This form is very rare here. Specimens prob¬ ably are smaller than Discorbinella valmonteensis Kleinpell (1938:350, pi. 21: figs. 14-16). It is described as “test large,” a point reiterated in the discussion. The holotype is at the National Mu¬ seum of Natural History. A/some paratype(s) and hypotypes reportedly were deposited in the Stan¬ ford collections but were not found. The small size of the Toro Road specimens is like that of the Valvulineria here. It may reflect environmental conditions. Yet, both specific and generic assignments are in question at this time. Literature Cited Barbat, W.F., and F.L. Johnson 1934. Stratigraphy and Foraminifera of the Reef Ridge Shale, Upper Miocene, California. Journal of Pa¬ leontology, 8:3-17. Buzas, M.A. 1965. The Distribution and Abundance of Foraminifera in Long Island Sound. Smithsonian Miscellaneous Collections, 149(1): 89 pages. 1968. On the Spatial Distribution of Foraminifera. Con¬ tributions from the Cushman Foundation for Fora- miniferal Research, 19: 11 pages. 1970. Spatial Homogeneity: Statistical Analyses of Un¬ ispecies and Multispecies Populations of Forami¬ nifera. Ecology, 51:874-879. 1978. Foraminifera as Prey for Benthic Deposit Feeders: Results of Predator Exclusion Experiments. Jour¬ nal of Marine Research, 36:617-625. Buzas, M.A., and T.G. Gibson 1969. Species Diversity: Benthonic Foraminifera in Western North Atlantic. Science, 163:72-75. Buzas, M.A., R.K. Smith, and K.A. Beem 1977. Ecology and Systematics of Foraminifera in Two Thalassia Habitats, Jamaica, West Indies. Smithson¬ ian Contributions to Paleobiology, 31: 139 pages. Buzas, M.A., C.F. Koch, S.J. Culver, and N.F. Sohl 1982. On the Distribution of Species Occurrence. Paleo¬ biology, 8:143-150. Byers, C.W. 1977. Biofacies Patterns in Euxinic Basins; a General Model. In Cook and Enos, editors, Deep-water Carbonate Environments. Society of Economic Pa¬ leontologists and Mineralogists, Special Publication, 25:5-18. Cushman, J. A. 1911. A Monograph of the Foraminifera of the North Pacific Ocean. United States National Museum, Bul¬ letin, 71(2): 108 pages. 1925. Some Textulariidae from the Miocene of Califor¬ nia. Cushman Laboratory for Foraminiferal Re¬ search, Contributions, l(2):29-35. 1926. Foraminifera of the Typical Monterey of Califor¬ nia. Cushman Laboratory for Foraminiferal Re¬ search, Contributions, 2(3):53-69. 1927. Recent Foraminifera from off the West Coast of America. Scripps Institution of Oceanography Tech¬ nical Series, Bulletin, 1:119—188. Cushman, J.A., and E.W. Galliher 1934. Additional New Foraminifera from the Miocene of California. Cushman Laboratory for Foramini¬ feral Research, Contributions, 10(l):24-26. d’Orbigny. See Orbigny, A. d’ Govean, F.M. 1980. Some Paleoecologic Aspects of the Monterey For¬ mation, California. 278 pages. Doctoral disserta¬ tion, University of California, Santa Cruz, Califor¬ nia. Govean, F.M., and R.E. Garrison 1981. Significance of Laminated and Massive Diatomites in the Upper Part of the Monterey Formation. In R.E. Garrison, R.G. Douglas, K.E. Pisciotto, C.M. Isaacs, and J.C. Ingle, editors, The Monterey For¬ mation and Related Siliceous Rocks, pages 181-198. Los Angeles: Pacific Section, Society of Economic Paleontologists and Mineralogists. Haller, C.R. 1980. Pliocene Biostratigraphy of California. In R.M. Kleinpell, editor, The Miocene Stratigraphy of Cali¬ fornia Revisited, pages 183-220. Tulsa: American Association of Petroleum Geologists. Ingle, G.C., G. Keller, and R.L. Kolpak 1980. Benthic Foraminiferal Biofacies, Sediments and Water Masses of the Southern Peru-Chile Trench Area. Micropaleontology. 26:113-150. Kleinpell, R.M. 1938. Miocene Stratigraphy of California. 450 pages, Tulsa: American Association of Petroleum Geol¬ ogists. Kleinpell, R.M., editor 1980. The Miocene Stratigraphy of California Revisited. 349 pages. Tulsa: American Association of Petro¬ leum Geologists. Kleinpell, R.M., and A. Tipton 1980. Taxonomy. In R.M. Kleinpell, editor, The Miocene Stratigraphy of California Revisited, pages 70-80. Tulsa: American Association of Petroleum Geol¬ ogists. Lankford, R.R. 1959. Distribution and Ecology of Foraminifera from East Mississippi Delta Margin. Bulletin of the Amer¬ ican Association of Petroleum Geologists, 43(9):2068-2099. Loeblich, A.R., Jr., and H. Tappan 1964. Sarcodina Chiefly “Thecamoebians” and Forami- niferida. In Raymond C. Moore, editor, Treatise on Invertebrate Paleontology, C(l-2): 900 pages. 27 28 Lawrence: University of Kansas Press for the Geo¬ logical Society of America. 1974. Recent Advances in the Classification of the For- aminiferida. In R.H. Hedley and C.G. Adams, editors, Foraminifera, 1:1-53. London and New York: Academic Press. Olsson, I., and B. Eriksson 1974. Horizontal Distribution of Meiofauna Within a Small Area, with Special Reference to Foramini¬ fera. Zoon, 2:67-84. Orbigny, A. d’ 1839. Foraminiferes. In de la Sagra, editor, Historie Phy¬ sique, Politique et Naturelle de Vile de Cuba, part 2 (Natural History), volume [7], 224 pages, unnum¬ bered plates. Paris: Bertrand. Parker, F.L., and W.D. Athearn 1959. Ecology of Marsh Foraminifera in Poponesset Bay, Massachusetts. Journal of Paleontology, 33:333-343. Phleger, F.B 1951. Ecology of Foraminifera, Northwest Gulf of Mex¬ ico, Part 1: Foraminifera Distribution. Geological Society of America Memoir, 46:1 -88. Phleger, F.B, and A. Soutar 1973. Production of Benthic Foraminifera in Three East Pacific Oxygen Minima. Micropaleontology, 19:1 10-115. Scott, G.H. 1958. Distribution of Populations of Fossil Foraminifera. SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY New Zealand Journal of Geology and Geophysics, l(3):474-484. Sen Gupta, B.K., R.F. Lee, and M.S. May, III 1981. Upwelling and an Unusual Assemblage of Benthic Foraminifera on the Northern Florida Continen¬ tal Slope. Journal of Paleontology, 55(4):853-857. Sheldon, A.L. 1969. Equitability Indices: Dependence on the Species Count. Ecology, 50:466, 467. Smith, R.K. 1970. Late Glacial Foraminifera from Southeast Alaska and British Columbia and a World-Wide High Northern Latitude Shallow-Water Faunal Prov¬ ince. Archives des Sciences, Geneve, 23(3):675-702. 1978. Systematics of the North American High North¬ ern Latitude Very Shallow Cold Water Forami- niferal Fauna. Archives des Sciences, Geneve, 31(2): 133—162. Stewart, R.E., and K.C. Stewart 1930. “Lower Pliocene” in Eastern End of Puente Hills, San Bernardino County, California. American As¬ sociation of Petroleum Geologists, Bulletin, 14:45- 50. Woodring, W.P., and M.N. Bramlette 1950. Geology and Paleontology of the Santa Maria District, California. United States Geological Suney Professional Paper, 222: 185 pages. PLATES 30 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY PLATE 1 Buliminella curta Cushman 1. Side view, hypotype, USNM 387632, X135. 2. Side view, hypotype, USNM 387633, X175. Buliminella elegantissima (d’Orbigny) 3. Side view, hypotype, USNM 387634, X180. Bolivina brevior Cushman 4. Side view, hypotype, USNM 387635, X250. 5. Apertural view, hypotype, USNM 387636, X165. 6. Side view, hypotype, USNM 387636, X155. 7. Apertural view, hypotype, USNM 387637, X260. 8. Side view, hypotype, USNM 387637, X220. Bolivina seminuda Cushman 9. Apertural view, hypotype, USNM 387638, Xl 25. 10. Side view, hypotype, USNM 387638, X140. 11. Side view, hypotype, USNM 387639, X95. 12. Side view, hypotype, USNM 387640, X205. Bolivina pseudospissa Kleinpell 13. Side view, hypotype, USNM 387641, X90. Suggrunda kleinpelli Bramlette 14. Apertural view, hypotype, USNM 387642, X190. 15. Combination view, hypotype, USNM 387642, X175. 16. Apertural view, hypotype, USNM 387643, Xl 55. 17. Side view, hypotype, USNM 387643, X190. NUMBER 60 31 32 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY PLATE 2 Bulimina cf. B. pseudoaffinis Kleinpell 1. Side view, hypotype, USNM 387644, X120. 2. Side view, hypotype, USNM 387645, X165. Globobulimina pacifica Cushman 3. Apertural view, hypotype, USNM 387646, x75. Epistominella subperuviana (Cushman) 4. Side view, hypotype, USNM 387647, X210. 5. Marginal view, hypotype, USNM 387647, X185. 6. Side view, hypotype, USNM 387648, X290. Valvulineria cf. V. californica obesa Cushman 7. Spiral view, hypotype, USNM 387649, X165. 8. Marginal view, hypotype, USNM 387650, X185. 9. Umbilical view, hypotype, USNM 387651, X160. 10. Spiral view, hypotype, USNM 387652, X200. 11. Marginal view, hypotype, USNM 387653, X215. 12. Umbilical view, hypotype, USNM 387654, X160. Nonionella schencki Kleinpell 13. 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