EBRIDIANOS FSILES EN LA LITERATURA CIENTFICA

RECOPILACIN DE DESCRIPCIONES E IMGENES (desde el ao 1973 al ao 2009)

Saunders, A.D., Larsen, H.C., and Wise, S.W., Jr. (Eds.), 1998 Proceedings of the Ocean Drilling Program, Scientific Results, Vol. 152

13. SILICEOUS SPONGE SPICULES, SILICOFLAGELLATES, AND EBRIDIANS FROM HOLE 918D, CONTINENTAL RISE OF THE GREENLAND MARGIN1
L. Kirk Lurvey,2,3 Kevin McCartney,2 and Wuchang Wei4

ABSTRACT Sediments recovered from Hole 918D on the continental rise of the Greenland Margin were found to contain siliceous sponge spicules, silicoflagellates, and ebridians. An extreme fossil-barren interval occurred in lower Miocene and upper Miocene sediments of the core, with sponge spicules occurring in the middle Miocene. Though generally rare, the sponge spicules were morphologically diverse, with monaxons being the most abundant. Other morphologies and an undescribed spicule type were found. Representatives of several silicoflagellate genera were found in a narrow horizon of the upper Pliocene. The silicoflagellates were of a variety of ages, showing that this layer has been reworked. This interval also included two large ebridians, Triskelion gorgon and Adonnadonna primadonna, that have not been previously described from the North Atlantic.

INTRODUCTION
Ocean Drilling Program (ODP) Leg 152 recovered sediment samples from six sites off the Greenland coast in order to study the rift volcanism and history of glaciation of the area. These sites were on a northwest-southeast transect across the East Greenland Margin, approximately 130 km from the Greenland coast near the center of a seaward-dipping reflector sequence (SDRS) (Fig. 1). A diverse assemblage of siliceous sponge spicules were found in middle Miocene sediments from Site 918D. The spicules were not generally abundant but were well preserved. The assemblage was dominated by monaxons with other spicule morphologies representing a fraction of total spicule abundance. The abundance and diversity of sponge spicules at Hole 918D are tabulated in this study (Table 1). Sponge spicules from deep ocean sediments have been studied infrequently. The results of this study can be compared to those of McCartney (1987, 1990), Ahlbach and McCartney (1992) and Zolnik et al., (1992); see McCartney (1990) for a tabulation of other sponge spicule literature from deep ocean studies. These studies show a generally similar diversity, with monaxons being the predominant spicule group. Palmer (1988) and Ivanik (1983) found abundant tetraxonal and triaxonal spicules and used their relative abundances to make paleobathymetric interpretations, but these spicule types were not abundant in this study.

Investigations were made using an Olympus BH-2 light microscope with a 20 objective. The entire coverslip was examined and absolute counts of specimens were recorded. In the event that no specimens were found on half of the coverslip, the rest was not investigated. In the narrow horizon where silicoflagellates and ebridians were present, extra slides were made and examined to increase specimen counts. The abundance of specimens was recorded in absolute counts, with pieces representing over half of the specimen, or enough to allow identification, counted as one.

SITE SUMMARY
Site 918D (635.572N, 3838.334W; water depth 1868.2 m) was drilled on the upper continental rise and has a thick Quaternary section. The site was selected to determine the age and subsidence history of the SDRS, the oceanographic history of the Irminger Basin, and the history of glaciation in southern Greenland. The core from Hole 918D provides the most complete sedimentary record recovered in the North Atlantic. Sponge spicules were found to be sparse in an 80-m interval of the middle Miocene and rare or absent elsewhere in the core. The sponge spicule-bearing interval is in lithologic Unit II, which is a silt with varying amounts of nannofossils and clay. The sponge spicules were always present in this interval but their numbers never exceeded 31 specimens in a slide. Silicoflagellates and ebridians were only found in two samples; all but one of the observed specimens came from Sample 152-918D13R-1, 9394 cm. This sample is from lithologic Subunit ID, which is a silt with dropstones.

METHODS
Raw samples were processed by disaggregating the sediment samples in 100-mL beakers after which H2O2 and 10% HCl solutions were added. The beakers were then heated for 34 hr on a hot plate. The remaining solutions were placed in test tubes and centrifuged and decanted three times. Smear slides were made from the remaining sediments using Canada balsam and 22 mm 50 mm coverslips.

SPONGE SPICULES
Sponges have an internal skeleton of spicules that may be of calcareous or siliceous composition. Various spicule shapes and sizes have been used to classify poriferans taxonomically (Ehrenberg, 1854). However, a single living sponge may contain a variety of spicule types. A descriptive terminology has been used to classify sponge spicules in recent literature (Bukry, 1978; Palmer, 1988; Ahlbach and McCartney, 1992), and is used in this study. Various types of sponge spicules are described here including monaxons, which have a skeletal element with a single axis; polyaxons, which have many equal-sized rays radiating from a single central point; and triaxons, which are spicules with three axes.

1 Saunders, A.D., Larsen, H.C., and Wise, S.W., Jr. (Eds.), 1998. Proc. ODP, Sci. Results,152: College Station, TX (Ocean Drilling Program). 2 Micropaleontology Undergraduate Research Laboratory, University of Maine at Presque Isle, Presque Isle, ME 04769, U.S.A. Correspondence author: McCartney@polaris.umpi.maine.edu 3 Department of Geology, University of Maine, Orono, ME 04469, U.S.A. 4 Scripps Institution of Oceanography, University of California at San Diego, La Jolla, CA 92093-0215, U.S.A.

191

L.K. LURVEY, K. MCCARTNEY, W. WEI Genus BACHMANNOCENA Locker, 1974; emend. Bukry, 1987 Bachmannocena apiculata (Schulz) (Pl. 3, Figs. 3, 4) Bachmannocena apiculata (Schulz), Bukry, 1987, p. 403404 Remarks. This silicoflagellate is common in sediments of the Eocene and Oligocene and suggests that the sediments in Sample 152-918D-13R-1, 92 94 cm have been reworked. Genus CORBISEMA Hanna, 1928 Corbisema hastata hastata (Lemmermann) (Pl. 3, Figs. 5, 6) Dictyocha triacantha hastata Lemmermann, 1901, p. 259, pl. 10, figs. 16, 17 Corbisema hastata hastata (Lemmermann), Bukry, 1976, p. 892, pl. 4, figs. 916 Remarks. The two specimens of this taxon were distinctly arrowhead shaped and were slightly asymmetrical. Corbisema triacantha (Lemmermann) (Pl. 3, Fig. 2) Corbisema apiculata (Lemmermann), Ling, 1972, p. 153, pl. 24, fig. 1 Remarks. The two specimens found of this taxon were slightly larger than typical C. triacantha. Genus DICTYOCHA Ehrenberg, 1837 Dictyocha messanensis Haeckel (Pl. 3, Fig. 10) Dictyocha messanensis Haeckel, in Peters (1860), p. 799800 Remarks. This taxon has considerable variability in the shape of its basal ring, the robustness of the skeletal elements, and in the presence or absence of an apical spine. Locker and Martini (1986) and other workers divided this taxon into several subspecies and forms, but these distinctions were difficult to apply consistently, thus, this taxon was not subdivided in this study. Genus DISTEPHANUS Stohr, 1880 Distephanus crux (Ehrenberg) (Pl. 3, Figs. 79) Distephanus crux crux (Ehrenberg), Bukry, 1976 [Leg 36], p. 895 Remarks. This was the most abundant silicoflagellate found in the study with a total count of 12. Several of these were variants with longer spines than usual for the species. A lone member of this species, shown on Plate 3, Fig. 9, was found in Sample 152-918D-39R-2, 9596 cm.; this was the only silicoflagellate in this study not found in Sample 152-918D-13R-1, 9294 cm. Genus NAVICULOPSIS Frenguelli, 1940 Naviculopsis constricta (Schulz) (Pl. 3, Fig. 1) Naviculopsis constricta (Schulz), McCartney and Wise, 1987, p. 807, pl. 5, figs. 58 Remarks. This problematic fossil has previously been found by Gombos (1982) in the middle Eocene of the southwest Atlantic Ocean and by McCartney and Wise (1990) in Oligocene sediments from the Weddell Sea near Antarctica. The occurrence in reworked sediments off Greenland suggests that this taxon is far more widely distributed, both geologically and geographically, than its sparse literature would indicate. Triskelion gorgon Gombos (Pl. 4, Figs. 24) Triskelion gorgon Gombos, 1982, p. 446447 Ebriopsis antiqua Schulz, 1928 (in part), p. 273274, fig. 696 Ebriopsis antiqua antiqua (Schulz), Ling, 1977, p. 215, pl. 1718 Genus TRISKELION Gombos, 1982 Ammodochium rectangulare (Schulz) (Pl. 4, Fig. 6) Ammodochium rectangulare (Schulz), Ling, 1971, p. 694 Genus CRANIOPSIS Hovasse ex Frenguelli, 1940 Craniopsis octo Hovasse ex Frenguelli, 1940 Craniopsis octo Hovasse ex Frenguelli, 1940, p. 92, figs. 31 ac Genus EBRIOPSIS Hovasse, 1932 Ebriopsis antiqua antiqua (Schulz) Adonnadonna primadonna (Gombos) (Pl. 4, Fig. 1) Adonnadonna primadonna Gombos, 1982, p.446 Remarks. This very unusual and large ebridian is similar and is probably closely related to Triskelion gorgon. Both have enormous size and exceptional surface ornamentation. For further remarks see those with Triskelion gorgon. Genus AMMODOCHIUM Hovasse, 1932

DISCUSSION
This study documents primarily the occurrence of sponge spicules in Hole 918D. Sample 152-918D-13R-1, 9394 cm, which lacked sponge spicules, was unusual in that it contained nearly all of the silicoflagellates and ebridians found in the study. The silicoflagellates and ebridians found in this sample were diverse, but include only one taxon, the silicoflagellate Dictyocha messanensis that is typically found in lower Pliocene sediments. The other taxa are reworked. Most of the reworked taxa are of geologically long-ranging species, making the age of the source material difficult to determine precisely. The presence of the ebridian Triskelion gorgon in the sample, however, indicates an age of middle Eocene to early Oligocene. All of the silicoflagellates found in this sample, except for the single specimen of Dictyocha messanensis, could be of the same age. The core description (Larsen, Saunders, Clift, et al., 1994) indicates that the sediment of Sample 152-918D-13R-1, 9394 cm was moderately disturbed. This is part of lithologic Unit ID, which consists of well-lithified quartz silt with clay and dark-gray compact sediment with isolated dropstones scattered throughout. The core description does not mention turbidites in this interval, although turbidites of two types are found in lithologic Unit IB, which occurred at 71.1236.0 mbsf. Several other studies of the Greenland transect also found reworking during the Pliocene or Pleistocene. Pliocene sediments at Site 919 included several foraminifers that were reworked (Israelson and

EBRIDIANS
Several representatives of ebridians were also found in Sample 152-918D-13R-1, 9294 cm. No ebridians were found elsewhere in the study.
Genus ADONNADONNA Gombos, 1982

194

SILICEOUS SPONGE SPICULES, SILICOFLAGELLATES, EBRIDIANS

Spezzaferri, this volume), and Wei (this volume) found Cretaceous nannofossils associated with dropstones in Pleistocene sediments, which he suggests were ice-rafted detritus. The reworked fossils of Sample 152-918D-13R-1, 9394 cm, might also be deposited as a result of ice-rafting, but it is more difficult to explain by this mechanism the concentration of typically uncommon and giant specimens such as Triskelion gorgon. Rafting should widely disperse the reworked specimens, and it would take unusual amounts of ice-rafting to deposit such a large concentration of silicoflagellates and ebridians in this sample. We believe that the concentration of siliceous microfossils in this sample can better be explained by a turbidity current.

CONCLUSIONS
The core from Hole 918D contained siliceous sponge spicules, silicoflagellates, and ebridians. The sponge spicules were found in the middle Miocene sediments, but were generally low in absolute abundance. The lower Miocene was largely barren, as was the upper Miocene and lower Pliocene. However, in the lower Pliocene there did occur an unusual presence of silicoflagellates and ebridians in Sample 152-918D-13R-1, 9394 cm. The core description indicates the sample was moderately reworked. This study found unusually large microfossils, including Triskelion gorgon, which suggests a turbidity current.

ACKNOWLEDGMENTS
We are grateful to Earl Oman for his slide preparation, help, and interest. We wish to thank Dr. Stuart Gelder for his assistance and technical advice. Sherwood W. Wise, Jr., and Andrew M. Gombos, Jr., provided valuable constructive comments on the manuscript. We thank the Ocean Drilling Program for providing the samples and the opportunity to conduct this study. Photographic material was contributed by the University of Maine at Presque Isle. This project, completed as an independent undergraduate research project, is Micropaleontology Undergraduate Research Lab publication No. 4.
REFERENCES Ahlbach, W.J., and McCartney, K., 1992. Siliceous sponge spicules from Site 748. In Wise, S.W., Jr., Schlich, R., et al., Proc. ODP, Sci. Results, 120: College Station, TX (Ocean Drilling Program), 833837. Bukry, D., 1976. Cenozoic silicoflagellate and coccolith stratigraphy, South Atlantic Ocean, Deep Sea Drilling Project Leg 36. In Hollister, C.D., Craddock, C., et al., Init. Repts. DSDP, 35: Washington (U.S. Govt. Printing Office), 885917. , 1978. Cenozoic coccolith, silicoflagellate, and diatom stratigraphy, Deep Sea Drilling Project Leg 44. In Bensen, W.E., Sheridan, R.E., et al., Init. Repts. DSDP, 44: Washington (U.S. Govt. Printing Office), 807863. , 1979. Cenozoic coccolith and silicoflagellate stratigraphy, Northern Mid-Atlantic Ridge and Reykjanes Ridge, Deep Sea Drilling Project Leg 49. In Luyendyk, B.P., Cann J.R., et al., Init. Repts. DSDP, 49: Washington (U.S. Govt. Printing Office), 551582. , 1987. Eocene siliceous and calcareous phytoplankton, Deep Sea Drilling Project Leg 95. In Poag, C.W., Watts, A.B., et al., Init. Repts. DSDP, 95: Washington (U.S. Govt. Printing Office), 395415.

Ehrenberg, C.G., 1854. Mikrogeologie: Das Erden und Felsen schaffende Wirken des unsichtbar kleines selbstndigen Lebens auf der Erde: Leipzig (Leopold Voss). Frenguelli, J., 1940. Consideraciones sobre los silicoflagelados fsiles. Rev. Mus. La Plata, Secc. Geol., 2:37112. Gombos, A.M., Jr., 1982. Three new and unusual genera of ebridians from the Southwest Atlantic Ocean. J. Paleontol., 56:444448. Hartman, W.D., 1982. Form and distribution of silica in sponges. In Simpson, T.L., and Volcani, B.E. (Eds.), Silicon and Siliceous Structures in Biological Systems: New York (Springer-Verlag), 453491. Ivanik, M.M., 1983. Paleogene and Neogene sponge spicules from Sites 511, 512, and 513 in the South Atlantic. In Ludwig, W.J., Krasheninnikov, V. A., et al., Init. Repts. DSDP, 71 (Pt. 2): Washington (U.S. Govt. Printing Office), 933950. Larsen, H.C., Saunders, A.D., Clift, P.D., et al., 1994. Proc. ODP, Init. Repts., 152: College Station, TX (Ocean Drilling Program). Lemmermann, E., 1901. Silicoflagellatae. Ber. Dtsch. Bot. Ges., 19:247271. Ling, H.Y., 1971. Silicoflagellates and ebridians from the Shinzan diatomaceous mudstone member of the Onnagawa Formation (Miocene), Northeast Japan. In Farinacci, A. (Ed.), Proc. 2nd Planktonic Conf. Roma. Rome (Ed. Technosci.), 689703. , 1972. Upper Cretaceous and Cenozoic silicoflagellates and ebridians. Bull Am. Paleontol., 62:135229. , 1977. Late Cenozoic silicoflagellates and ebridians from the eastern North Pacific region. In Saito, T., and Ujiie, H. (Eds.), Proc. First Int. Congr. Pacific Neogene Stratigraphy, 1:205233. Locker, S., and Martini, E., 1986. Silicoflagellates and some sponge spicules from the southwest Pacific, DSDP Leg 90. In Kennett, J.P., von der Borch, C.C., et al., Init. Repts. DSDP, 90: Washington (U.S. Govt. Printing Office), 887924. McCartney, K., 1987. Siliceous sponge spicules from Deep Sea Drilling Project Leg 93. In van Hinte, J.E., Wise, S.W., Jr., et al., Init. Repts. DSDP, 93 (Pt. 2): Washington (U.S. Govt. Printing Office), 815824. , 1990. Siliceous sponge spicules from Ocean Drilling Program Leg 113. In Barker, P.F., Kennett, J.P., et al., Proc. ODP, Sci. Results, 113: College Station, TX (Ocean Drilling Program), 963970. McCartney, K., and Wise, S.W., Jr., 1987. Silicoflagellates and ebridians from the New Jersey Transect, Deep Sea Drilling Project Leg 93, Sites 604 and 605. In van Hinte, J.E., Wise, S.W., Jr., et al., Init. Repts. DSDP, 93 (Pt. 2): Washington (U.S. Govt. Printing Office), 801814. , 1990. Cenozoic silicoflagellates and ebridians from ODP Leg 113: biostratigraphy and notes on morphologic variability. In Barker, P.F., Kennett, J.P., et al., Proc. ODP, Sci. Results, 113: College Station, TX (Ocean Drilling Program), 729760. Palmer, A., 1988. Paleoenvironmental significance of siliceous sponge spicules from Sites 627 and 628, Little Bahama Bank, Ocean Drilling Program Leg 101. In Austin, J.A., Jr., Schlager, W., et al., Proc. ODP, Sci. Results, 101: College Station, TX (Ocean Drilling Program), 159168. Peters, W., 1860. Kurzen Auszug aus einer Abhandlung des Hrn. Dr. Ernst Haeckel ber neue, ledende Radiolarien des Mittelmeeres und legte die dazu gehrigen Abbildungen vor. Mber. Verh. K. Preuss. Akad. Wiss. Berlin, 794817. Schulz, P., 1928. Beitrge zur Kenntnis fossiler und rezenter Silicoflagellaten. Bot. Arch., 21:225292. Zolnik, R., McCartney, K., and White, L.D., 1992. Siliceous sponge spicules from Site 795. In Pisciotto, K.A., Ingle, J.C., Jr., von Breymann, M.T., Barron, J., et al., Proc. ODP, Sci. Results, 127/128 (Pt. 1): College Station, TX (Ocean Drilling Program), 541544.

Date of initial receipt: 14 August 1995 Date of acceptance: 12 June 1996 Ms 152SR-216

195

SILICEOUS SPONGE SPICULES, SILICOFLAGELLATES, EBRIDIANS

Plate 4. Ebridians (570; scale bar = 35 m). 1. Adonnadonna primadonna Gombos; Sample 152-918D-13R-1, 9394 cm. 24. Triskelion gorgon Gombos; Sample 152-918D-13R-1, 9394 cm. 5. Unknown ebridian, Sample 152-918D-13R-1, 9394 cm. 6. Ammodochium rectangulare (Schulz), lorica; Sample 152918D-13R-1, 9394 cm.

199

Tracking Environmental Change Using Lake Sediments Volume 3: Terrestrial, Algal, and Siliceous Indicators
Edited by

John P. Smol
Department of Biology, Queens University

H. John B. Birks
Botanical Institute, University of Bergen
and

William M. Last
Department of Geological Sciences, University of Manitoba

KLUWER ACADEMIC PUBLISHERS

NEW YORK, BOSTON, DORDRECHT, LONDON, MOSCOW

10. EBRIDIANS

ATTE KORHOLA (atte.korhola@helsinki.fi)

Department of Ecology and Systematics Division of Hydrobiology University of Helsinki P. O. Box 17 (Arkadiankatu 7) FIN-00014 Helsinki, Finland
JOHN P. SMOL (SmolJ@BIOLOGY.QueensU.Ca)

Paleoecological Environmental Assessment and Research Lab (PEARL) Department of Biology 116 Barrie St., Queens University Kingston Ontario K7L 3N6 Canada
Keywords: Ebridians, Ebria tripartita, sediment, Baltic Sea, palaeoecology, eutrophication, dissolution

Introduction

Ebridians are a group of microscopic, heterotrophic, primarily marine plankton, characterized by a siliceous internal skeleton that is frequently preserved in sedimentary deposits. They are cosmopolitan, mainly neritic, but can also be found in estuaries, shallow embayments, and semi-enclosed seas such as the Baltic Sea and the Black Sea (Ernisse & McCartney, 1993; Korhola & Grnlund, 1999). Ebridians have a modern diversity of only a few species, and are not particularly significant components of modern marine plankton. However, in contrast to the diatoms, where perhaps 90% of the described species are living, most of the described ebridian taxa are fossil forms. Indeed, ebridian research is essentially palaeontologic, and they are widely used in stratigraphy in palaeoceanography and in palaeobiologic studies. As such, the connection of ebridians to a volume on palaeolimnological methods and indicators may be a bit tenuous, but they are occasionally recorded in freshwater (likely from allochthonous sources) and brackish profiles, and so this short chapter attempts to provide a brief overview of this group which may be of use to the palaeolimnologist. Our primary aim is to attract attention to these organisms, whose remains are found regularly in the sediment record, but are often overlooked. In addition, we explore the potential usefulness and indicator value of the group in future palaeoecological studies.
225 J. P. Smol, H. J. B. Birks & W. M. Last (eds. ), 2001. Tracking Environmental Change Using Lake Sediments. Volume 3: Terrestrial, Algal, and Siliceous Indicators. Kluwer Academic Publishers, Dordrecht, The Netherlands.

226

ATTE KORHOLA & JOHN P. SMOL

We focus in particular on one species, Ebria tripartita (Schumann) Lemmermann, and one environment, namely the brackish-water Baltic Sea, where this species has been studied quite intensively during recent years. Ernisse & McCartney (1995) provide a more detailed summary of these indicators, and Loeblich et al. (1968) provide an annotated index of ebridian taxa.

Morphology, taxonomy and preservation in the sediments Ebridians do not appear in the fossil record before the Cenozoic, but their fossils have been found in particularly large numbers in Cenozoic sedimentary rocks that are rich in various other siliceous remains (Tappan, 1980). After a phase of rapid diversification in the Paleocene, during which at least eight ebridian genera evolved, they suffered massive extinctions by the close of the Miocene. Although they have a relatively diverse fossil history, ebridians are relatively rare in the sediments of modern seas. Only three (possibly four) species, assigned to two genera (Ebria and Hermesinum), are known to exist today (Taylor, 1990). The ecological characteristics of the few living ebridians are poorly known, and so they have played a relatively small role in modern research. Ebridians commonly range from 10 to in length and from 20 to in width (Lipps, 1979). Their cells are more or less spherical, colourless, and contain a single nucleus and two flagella of different lengths (Lipps, 1979; Tikkanen, 1986). Some ebridian taxa produce cyst-like features at one end of the skeleton that may have served as a protective enclosure during periods of environmental stress (Ernisse & McCartney, 1995). Ebridians are preserved in bottom deposits because of their internal siliceous skeleton, which consists of a framework of solid rods, in thickness (Preisig, 1994). In Ebria tripartita, the system of rods comprises three stems, which branch in a regular manner from an initial branching point, thus forming a triadial, basket-like symmetry (Figs. 1 and 2). In Hermesinum four such branches are found. The rods are usually angular and possess many small spines and nodules, as shown by electron microscopy (Fig. 2B). The average length of the siliceous skeleton of E. tripartita is and the average width is (Tikkanen, 1986). The siliceous skeleton of ebridians is highly resistant both against mechanical abrasion and chemical dissolution. The taxonomic status of ebridians is presently rather uncertain (Preisig, 1994). On one hand, they resemble silicoflagellates, but have a skeleton of solid tubular rods rather than hollow ones. On the other hand, the structure of their nuclei as well as their endoskeleton has some common features with some dinoflagellates (Loeblich, 1982; Lipps, 1979; Locker & Martini, 1986). Indeed, the evolutionary relations among ebridians, silicoflagellates, and endoskeletal dinoflagellates are in many respects unclear. Today, the ebridians are, nevertheless, often regarded as a separate taxonomic unit (Ebriophyceae/Ebriida) with uncertain phylogenetic affinities (Taylor, 1990; Preisig, 1994). In the older literature, there have been erroneous identifications concerning these taxa. Most common mistakes include the assignment of the species with other dinoflagellates, silicoflagellates or even Radiolaria. It is perhaps because of these identification problems that these taxa have rarely been recognized and have only been used in a few palaeolimnological studies.

EBRIDIANS

227

Methodological aspects The chemical treatment of subsamples follows the standard methodology developed for diatom sample preparation (Battarbee, 1986 and Battarbee et al., this volume). Naphrax or Hyrax or comparable media can be used as mountants. Counting is performed under a light microscope at 1000 magnification. The frequencies of ebridians can be determined relative to diatoms (Korhola & Grnlund, 1999) or the total siliceous microfossil assemblage (Westman, 1999), which can then be based on the sum of total diatoms/siliceous fossils + ebridians. In general, ebridians tend to be rather rare, occurring in low absolute frequencies in an average microfossil preparation (Ernisse & McCartney, 1995). Moreover, their remains are often fragmented (Korhola & Grnlund, 1999). The preservation of the main part of the central skeleton (a complete driadial system of rods) can be used as the criterion for the enumeration of one species. Brief history of use of ebridians in palaeoecological research Ebridian research has developed episodically, with an initial phase of interest in the nineteenth century, a quiet interval between about 1910 and 1950, and a second, mostly geologic phase of development since then, in conjunction with renewed interest in subjects such as plate tectonics and palaeoceanography after WWII. Most extensive and reliable data about the biostratigraphic ranges of ancient ebridian species have been generated during the Deep Sea Drilling Project (DSDP, 1968 to 1983) and the Ocean Drilling Project (ODP, 1985 to present). For example, Locker & Martini (1989) established nine ebridian zones that span the interval from the Early Miocene to the early Quaternary, and demonstrated that these organisms might have some value as biostratigraphic markers and environmental indicators. The work dealing with ancient ebridians (i. e., pre-Quaternary biostratigraphy) is referenced and discussed in more detail in Ernisse & McCartney (1995), whereas here

228

AITE KORHOLA & JOHN P. SMOL

we discuss the few living ebridians and their applicability to more recent environmental and palaeolimnological problems. Little is known about the present-day ecological preferences of the ebridians, which, of course, hinders palaeoecological interpretations. According to Lipps (1979), they are very abundant in nutrient-rich, cool marine waters at high latitudes. It seems that from the two main living species, Hermesinun adiraticum is a stenohaline organism that thrives in warm waters, whereas E. tripartita has a wider tolerance to temperature and salinity variations (Rhodes & Gibson, 1981). The latter is the only ebridian species alive in the Baltic Sea today, and intact remains of E. tripartita, are frequently found in sediments representing the ancient history of the Baltic basin, such as the Eemian Baltic Sea about 120 000 yrs BP (Brander, 1937; Niemel & Tynni, 1979; Eriksson et al., 1980; Grnlund, 1991). In the Baltic Sea, increased abundances of this taxon are regularly noted in the uppermost parts of sediment cores and are commonly interpreted as indicating nutrient enrichment and/or upwelling (e. g., Miller & Risberg, 1990; Grnlund, 1991, 1993; Saxon & Miller, 1993; Andrn, 1995). For example, Miller & Risberg (1990) noted increased abundances of Ebria tripartita in surface sediments of a core from the northwestern Baltic, and related this change to the acceleration of eutrophication during the last ca. 20 years. Similarly, Korhola & Gronlund (1999) observed in the Gotland basin core a slight increase in the abundance of E. tripartita towards the core surface. This increase was also tentatively interpreted as being related to the progressive eutrophication of the basin. Numerous studies (e. g., Rosenberg,

EBRIDIANS

229

1984; Elmgren, 1989; Baltic Marine Environment Protection Commission, 1990) point to a marked increase in nutrients in the Baltic Sea during the last 200 years. However, in the sediment core from Tl Bay (Fig. 3), central Helsinki (Finland), the abundances of E. tripartita were particularly low during the most pronounced eutrophication stage, as inferred from diatoms (Korhola & Blom, 1996; Korhola & Grnlund, 1999). One explanation could be that, despite the species general preference for nutrient-rich waters, it might be less competitive in hypereutrophic waters, especially if the eutrophication is associated with heavy blooms of other planktonic algae (Korhola & Grnlund, 1999). On the other hand, studies of recent blooms in Long Island Sound USA (Ernisse & McCartney, 1993) and the Baltic Sea (Kononen & Niemi, 1984) have observed significant year-to-year fluctuations in the absolute abundance of ebridians, which do not seem to be related to changes in either salinity or trophic status. In recent years, there has been an increased interest in modern (extant) forms found as fossils in the surface sediment of the shallow seas. Westman (2000) studied four long sediment cores, five short cores, and more than 20 surface-sediment samples representing the years 1993 and 1997 for subfossil ebridians in die Baltic Sea proper. He was unable to distinguish any systematic trend in the abundance of E. tripartita during the most recent centuries, although a slight increase in the relative abundance of this taxon was observed in all sub-surface cores. All variation in the ebridian data seemed to be confined to periods

230

ATTE KORHOLA & JOHN P. SMOL

when there were also major changes in diatom assemblagesa situation that seems to hold true in other cases as well (e. g., Korhola & Grnlund, 1999). This prompted Westman (1999) to conclude that the changes in the abundance of E. tripartita are "most probably attributable to alterations in the species composition of primary production in the Baltic, and hence represent only a secondary effect of environmental change". As E. tripartita mainly feeds on diatoms (Ernisse & McCartney, 1993), changes in its relative frequency could simply result from changes in the composition of the diatom flora and/or the contribution of the diatom flora to total primary production (Westman, 1999). This feature is illustrated in Figure 4, where the abundances of E. tripartita in the surface-sediments of the Baltic Sea proper sampled in 1993 and 1997 are contrasted against the major differences in diatom assemblages, as revealed by cluster analysis (Westman, 1999). As the abundance of E. tripartita in the sediments seems to be reflected in major differencies in the diatom assemblage, it may be assumed that the distribution of the species in the sediments is predominantly controlled by differencies in the composition of the diatom assemblage. In an ongoing study dealing with the eutrophication history of the shallow Baltic Sea inlets, ebridians have been identified together with diatoms in 45 sampling stations along the coastal area of southern Finland (Weckstrm & Korhola, in press). Only a few remains

EBRIDIANS

231

of E. tripartita were found, with their absolute numbers ranging between 012 per 500 diatom frustules in sediment samples. A preliminary principal components analysis (PCA) performed on the measured water chemistry and ebridian data suggests that the ebridian distributions and abundances in this data set are correlated with the PCA axis 1 consisting of salinty and ionic concentration (Fig. 5). This axis explained 46. 6% of the variance in the environmental and ebridian data. It thus seems that ebridians in these shallow embayments prefer sites of relatively high salinity (>4psu), conductivity and, to some extent, concentrations. Further studies will focus on the relationships of ebridians in these shallow coastal waters to various diatom species and assemblages. Indicator value and future research priorities At present, contradictory data regarding the indicator value of ebridians exist, which are most obviously related to the rudimentary ecological data available for these organisms.

232

ATTE KORHOLA & JOHN P. SMOL

As shown by the examples discussed in this chapter, ebridians clearly show stratigraphic changes in sediment profiles, yet the ecological interpretations of these changes are still speculative. Clearly, more systematic studies on the distribution of these organisms along a range of various ecological gradients are needed in order to achieve a more comprehensive understanding of the palaeolimnological significance of the group. Future studies should focus on the collection of modern surface-sediment data sets along a wide range of environments and ecological conditions in different marine systems. Some results indicate that the observed patterns in the frequency distributions of ebridians may be partly due to the selective preservation of their siliceous skeletons (Korhola & Grnlund, 1999). Thus, the questions dealing with the preservation and taphonomy of these organisms should be researched more fully. The taxonomy of ebridians is in many respects vague; thus in order for ebridians to become more useful indicators, detailed taxonomic studies using both light and electron microscopy are needed. Moreover, if ebridians are strongly affected by grazing pressure, as suggested by Westman (1999), then the question of which food web changes might affect ebridians would be an interesting research question. Summary Ebridians are a group of microscopic, heterotrophic marine plankters. Their siliceous endoskeletons are preserved in sedimentary deposits, and so they can be studied using the same techniques developed for other siliceous indicators, such as diatoms. They are primarily marine, and so are not frequently encountered by palaeolimnologists, but they may be common in brackish waters, such as the Baltic Sea, estuaries, and some lacustrine environments that may have had an influx of marine material. Only a few species are known to exist today. Relatively little is known about the ecological optima and tolerances of taxa, which currently hampers palaeoecological interpretations. However, ongoing research suggests they have some potential in palaeoenvironmental studies. Acknowledgements The study was funded by the Academy of Finland Grant 101 7383 to AK. We thank Kaarina Weckstrm for counting the ebridian remains from the surface-sediments of the Gulf of Finland. References
Andrn, E., 1995. Recording environmental changes in the Southern Baltic Seacurrent results from a diatom study within Project ODER. In Marino, D. & M. Montesor (eds. ) Proceedings of the 13th International Diatom Symposium. Maratea, Italy, lst-7th September 1994: 443455. Baltic Marine Environment Protection Commission, Helsinki Commission, 1990. Second Periodic Assessment of the State of the Marine Environment of the Baltic Sea, 19841988; Background Document. Baltic Sea Environment Proceedings 35 B, 432 pp. Brander, G., 1937. Zur Deutung der intramornen Tonablagerungen an der Mga. unweit von Leningrad. Bull. Comm. Gol. Fin. 119: 93113. Edler, L., G. Hllfors & . Niemi, 1984. A preliminary check-list of the phytoplankton of the Baltic Sea. Acta Bot. Fenn. 128, 26 pp.

EBRIDIANS

233

Elmgren, R., 1989. Mans impact on the ecosystem od the Baltic Sea: Energy flows today and at the turn of the century. Ambio 6: 326332. Eriksson, B., T. Grnlund & R. Kujansuu, 1980. Interglasiaalikerrostuma Evijrvell Pohjanmaalla (English summary: An interglacial deposit at Evijrvi in the Pohjanmaa region). Geologi 32: 6571. Ernisse, J. & K. McCartney, 1993. Ebridians. In Lipps, J. H. (ed. ) Fossil Prokaryotes and Protists. Blackwell Scientific Publication Inc., Boston: 131140. Ernisse, J. & McCartney, 1995. Ebridians and endoskeleton dinoflagellates. In Blome, C. D., P. Whalen & K. Reedet (eds. ) Siliceous Microfossils. Paleontology Society Short Courses in Paleontology 8: 177185. Grimm, E., 1987. CONISS: A Fortran 77 program for stratigraphically constrained cluster analysis by the method of incremental sum of squares. Comp. Geosci. 13: 1325. Gronlund, T., 1991. The diatom stratigraphy of the Eemian Baltic Sea on the basis of sediment discoveries in Ostrobothnia. Finland. Geol. Surv. Fin., Report 102, 26 pp. Grnlund, T., 1993. Diatoms in surface sediments of the Gotland Basin in the Baltic Sea. Hydrobiologia 269/270: 235242. Hakansson, H. & . Korhola, 1998. Phenotypic plasticity in the diatom Cyclotella meneghiniana or a new species? Nova Hedwigia 66: 187196. Kononen, K. & . Niemi, 1984. Long term variation of the phytoplankton composition at the entrance to the Gulf of Finland. Ophelia suppl. 3: 101110. Korhola, A. & T. Blom, 1996. Marked early 20th century pollution and the subsequent recovery of Tl Bay, central Helsinki, as indicated by subfossil diatom assemblage changes. Hydrobiologia 341: 169179. Korhola, A. & T. Grnlund, 1999. Observations of Ebria tripartita (Schumann) Lemmermann in Baltic sediments. J. Paleolim. 21: 18. Lipps, J., 1979. Ebridians. In Fairbridge, R. W. & D. Jablonski (eds. ) Encyclopedia of Earth Sciences. Vol 7. Dowden, Hutchinson & Ross, Stroudsburg, 276 pp. Locker, S. & E. Martini, 1986. Ebridians and actiniscidians from the southwest Pacific. In Kennett, J. P., et al. (eds. ) Initial Reports of the Deep Sea Drilling Project. 90. U. S. Goverment Printing Office, Washington, D. C.: 939951. Locker, S. & E. Martini, 1989. Cenozoic silicoflagellates, ebridians, and actiniscidians from the Vring Plateau (ODP Leg 104). In Eldholm, O., et al. (eds. ) Proceedings of the Ocean Drilling Program, Scientific results. 104. Ocean Drilling Program, College Station, Texas: 543585. Loeblich, A. R. III, 1982. Dinophyceae. In Parker, S. P. (ed. ) Synopsis and Classification of Living Organisms. Vol 1. McGraw-Hill, New York: 101115. Loeblich, A. R. III, A. L., Loeblich, H., Tappan & A. R. Loeblich, Jr., 1968. Annotated index of fossil and recent silicoflagellates and ebridians with descriptions and illustrations of validly proposed taxa. Geological Society of America, Memoire 106: 1319. Miller, U. & J. Risberg, 1990. Environmental changes, mainly eutrophication, as recorded by fossil siliceous micro-algae in two cores from the uppermost sediments of the north-western Baltic. Nova Hedwigia 100: 237352. Niemel, J. & R. Tynni, 1979. Interglacial and interstadial sediments in the Pohjanmaa region. Finland. Geol. Surv. Fin. Bull. 302, 48 pp. Preisig, H. R., 1994. Siliceous structures and silicification if flagellated protists. Protoplasma 181: 2942. Rhodes, R. G. & V. R., Gibson, 1981. An annual survey of Hermesinum adriaticum and Ebria tripartita, two ebridian algae in the lower Chesepeake Bay. Estuaries 4: 150152. Rosenberg, R. (ed. ), 1984. Eutrophication in Marine Waters Surrounding Sweden, A review. Statens naturvrdsverk, Solna. PM 1808, 140 pp.

234

ATTE KORHOLA & JOHN P. SMOL

Saxon, M. & U. Miller, 1993. Diatom assemblages in superficial sediments from the Gulf of Riga, eastern Baltic Sea. Hydrobiologia 269/270: 243249. Tappan, H., 1980. The Paleobiology of Plant Protists. Freeman, San Francisco, 1028 pp. Taylor, F. J. R., 1990. Ebridians. In Margulis, L., J. O. Corliss, M. Melkonian & D. J. Chapman (eds. ) Handbook of Protoctista. Jones and Bartlett, Boston, 720721. Tikkanen, T, 1986. Kasviplanktonopas (Guide to Phytoplankton). Suomen Luonnonsuojelun Tuki Oy, Forssa, 278 pp. Weckstrm, K. & A. Korhola. Physical and chemical characteristics of 45 shallow embayments on the southern coast of Finland. Hydrobiologia, in press. Westman, P., 2000. The Siliceous microalgae Dictyocha speculum and Ebria tripartita as biomarkers and palaeoecological indicators in Holocene Baltic Sea sediments. GFF 112: 287292.

Hydrobiologia 473: 217221, 2002. D.A. Barnum, J.F. Elder, D. Stephens & M. Friend (eds), The Salton Sea. 2002 Kluwer Academic Publishers. Printed in the Netherlands.

217

Skeletal development in Hermesinum adriaticum Zacharias, a agellate from the Salton Sea, California
M. A. Tiffany
Department of Biology, San Diego State University, U.S.A. Key words: Ebriidians, phytoplankton, salt lake

Abstract Hermesinum adriaticum is a rarely reported unicellular biagellated organism with a solid siliceous skeleton. Live specimens were not observed but many skeletons were found in sediments and, in small numbers, in cyanobacterial mats and the water column of the Salton Sea, a salt lake in California, U.S.A. Stages of the developing skeleton were studied with scanning electron microscopy, and the progression from small tetraxial daughter skeletons to complete asymmetrical adult skeletons is presented. Some variability in the adult skeletons is illustrated. Introduction Ebriidians are a rare group of marine heterotrophic agellates of uncertain classication. Corliss (1994) has tentatively placed them in his phylum Opalozoa, and Taylor (1990) put them as possibly related to the dinoagellates. Ebriidians have two dissimilar agella and solid internal siliceous skeletons in contrast to the external hollow skeletons of silicoagellates. There are two extant genera validly described, Ebria Borgert and Hermesinum Zacharias, of these Ebria is the more commonly reported in plankton surveys. Presently, H.adriaticum Zacharias and H. platense Frenguelli are the only two known extant species in the genus. Others have been recorded as fossils, mostly from the Miocene (Tappan, 1980). The type of H. adriaticum was from the Adriatic Sea (Zacharias, 1906). Occasional blooms of H. adriaticum have been recorded from the southern Mediterranean near the Nile River (Halim, 1960), the Black Sea (Bodeanu, 1969), the Pettaquamscutt River in Rhode Island (Hargraves & Miller, 1974), the lower Chesapeake Bay (Rhodes & Gibson 1981) and Lake Rogoznica, a salt lake near the eastern Adriatic coast (Vilicic et al., 1996/97). The greatest abundance of H.adriaticum found to date was in the Salton Sea, California in 1955 (Carpelan, 1961) at 450 cells/ml, although it was misidentied as a silicoagellate, possibly Dictyocha sp.?. For a photograph of a living cell of H. adriaticum, see Rhodes & Gibson (1981).

Figure 1. Diagram showing the terminology for parts of the Hermesinum adriaticum skeleton (after Deandre, 1952).

Reproduction of the ebriidian cell has been little studied, and only asexual division is known (Tappan, 1980). Formation of the skeleton of the daughter cell is thought to begin before nuclear division and be nearly complete before cellular division. Rarely, double skeletons are formed by the accidental fusion of the daughter with the original skeleton (Hovasse, 1932). Skeletal parts are named using the traditional nomenclature of sponge spicules as proposed by Deandre (1952) and are shown in Figure 1.

218 Study area The Salton Sea is a saline lake in Imperial County, California that is below sea level and receives inows from agricultural and municipal wastewaters. It began as a freshwater lake formed as a result of an accidental diversion of the Colorado River in 1905. Due to the lack of an outow and the evaporation rates in its desert environment, the salinity rose gradually. When Carpelan (1961) reported high densities of H. adriaticum in 19551956, the lake had a salinity close to that of seawater. Presently it has a salinity of approximately 43 g l1 . Scanning electron microscope images of adult skeletons of H.adriaticum from the Salton Sea, taken by A. R. Loeblich from 1967 to 1969, have previously been published (Tappan, 1980). Marine micro-organisms were likely introduced accidentally along with marine sh that were stocked into the Salton Sea, mostly in the 1940s and 1950s. Many of these sh were from the Gulf of California, some from the Pacic Ocean and coastal Texas. Most of the organisms in the Sea are of marine origin,including H. adriaticum, possibly as a direct result of these introductions. Migratory birds are a less likely source of inoculation because the life cycle is not known to include a cyst form (Tappan, 1980). H. adriaticum has an optimum temperature of 20 25 C (Hargraves & Miller, 1974), well within the range for the Salton Sea of 1235 C. The salinity tolerance is unknown, but it has been reported at salinities as low as about 12 g l1 (Hargraves & Miller 1974) and as high as 38 g l1 (Vilicic et al., 1996/97). Samples of oating cyanobacterial mats were taken from Varner Harbor at the headquarters of the Salton Sea State Recreational Area. A mid-lake water sample was taken by bucket from the surface (approximately the top 10 cm) from the northern basin (33 23 23.9 N, 115 51 57.0 W) on 4/7/99. All samples were treated by the Van Stosch method (Hasle & Syvertsen, 1997) using HNO3 and H2 SO4 to remove organic material, dried on a coverslip, coated with Au/Pd, and examined with a Hitachi 5200 SEM using 10 KV accelerating voltage.

Results and discussion H. adriaticum, previously known to be present in the Salton Sea, (Carpelan, 1961; Tappan, 1980) was found in several types of samples in 19981999. The largest numbers were found in the sediment sample. The sediment sample contained a great many individual skeletons of H.adriaticum. Both adult and daughter skeletons were found. The number of skeletons in the surcial sediments suggests a bloom occurred in the recent past. Several adult and daughter skeletons from organisms that may have been alive when collected were found on SEM stubs of samples of cyanobacterial mats that had been processed with concentrated acids. These mats rise to the surface in summer and contain many entrained benthic diatoms as well as the much rarer H.adriaticum. H. adriaticum is thought to be herbivorous on diatoms (Tappan, 1980) or agellates (Hargraves & Miller, 1974). A single adult skeleton was found on a stub prepared from a mid-lake surface water sample. This may represent resuspension from the sediment or a living specimen at the time it was collected. No living specimens of H. adriaticum were observed during this study. A complete progression of daughter skeletons was found in samples from small tetraxial daughter skeletons about 10 m long to adult skeletons 3754 m long (mean=44.4 m, n=29). The young skeletons, called trianes, have four rays called the actines. Three of these are set at about 120 apart in a plane, the fourth projects at a 90 angle from that plane (Fig. 2A, B). The fourth actine is topped by a triangle of silica (Fig. 2B, C). The two anterior apices of the triangle grow extensions that close to become the small circular upper window (Fig. 2D F). The posterior apex of the triangle lengthens but

Materials and methods A study of diatoms was carried out as part of a larger bio-inventory of the Salton Sea during 1999. Skeletons of H. adriaticum were encountered during the SEM analysis of samples prepared for the determination of diatom species. Surface sediment was sampled on January 20, 1999 from mid-lake stations by L. F. R. LevineFricke as part of a study of sediment contaminants using a modied Birge-Eckman dredge. One of these samples (GB64-46.2-12099), taken at 33 23 23.9 N, 115 51 57.0 W (about 4 km from the eastern shore in the northern basin) was donated to the author for study of diatoms and other microfossils. Depth at the site was 13 m.

219

Figure 2. Stages in the development of and variability in skeletons of Hermesinum adriaticum. (A and B) very young skeletons showing the primary four-sided symmetry. (C and D) beginning of secondary development of the apices of the original skeleton. (E) small dorsal window beginning to form and continuation of apical growth with large upper window now fully formed. (F) adult left-handed skeleton with fully formed opisthoclade and spinose surface. (G) an adult right-handed skeleton showing smoother surface than the individual in F. (H) Error in skeleton formation with misplaced anterior apex and an extension to dorsal apex instead of an opisthoclade. (I and J) Double anterior apices. (K) illustration of rugose texture of skeleton at higher magnication. (L) spiny surface of adult skeleton. Scale bar for AJ=10 m, for K=1 m, and for L=5 m.

normally does not connect further to the skeleton. The three bottom actines extend during development, the two anterior ones trifurcating (Fig. 2C). The third,

posterior actine becomes the longer rhabde, forming the axis of the skeleton. It has three extensions, one of which becomes the posterior spine and one that

220 will help form the opisthoclade, an arch-like feature. The anterior extensions extend in a curved manner and fuse to form the large upper window (Fig. 2E). The anterior spine is formed at the forward end of this large window. The posterior extension of one of the anterior actines fuses with that of the rhabde to make the opisthoclade in the adult skeleton (Fig. 2E, F). The posterior extension of the other anterior actine forms the lateral spine. The small upper window forms two bridges on either side to the lower anterior window forming a sort of siliceous basket in which the nucleus of the daughter cell will reside (Fig. 2E). At which point in the skeletal development the nucleus divides and exactly when the daughter nucleus enters the newly forming skeleton is not known, but Deandre (1952) illustrated a cell undergoing cytokinesis with an incomplete daughter skeleton, the nuclear basket of silicon not yet formed. Skeletons of Hermesinum are always asymmetric with a completed arch only on one side of the skeleton. Assuming that Figure 1 illustrates the dorsal view of the skeleton (after Hargraves & Miller, 1974), the right-handed skeleton is dened as one where the opisthoclade is on the right side of the skeleton. The illustration of a double skeleton given by Hovasse (1932) shows fused daughter skeletons that are mirror images. This suggests that after division one daughter is left-handed (Fig. 2F) and one is right-handed (Fig. 2G). This would imply that in a population of this species equal numbers of left and right-handed skeletons should be present. Of the 29 adult skeletons of Hermesinum examined, 15 had the arch on the left side and 14 on the right, in close agreement with this hypothesis. Much variability in the development of the skeleton occurs in the ebriidians. The morphology of several skeletons in this study differed substantially from the diagram in Figure 1. For example, in Figure 2H the upper triangle of silica apparently extended to the posterior apex, forming a bridge; an error observed in several specimens. In Figure 2I, there are two small upper windows instead of one and in Figure 2J two anterior apices were accidentally formed. In these samples, no double skeletons were observed but they are known to be common in the ebriidians (Tappan, 1980). It is not clear what, if any, effect this variability might have on the cell. At high magnication, the texture of the skeleton is evident (Fig. 2K). In contrast to the relative smoothness seen in some silica secreting organisms, such as silicoagellates and some diatoms, it is rather rugose. The skeleton is solid and internal to the cell. This is in contrast to silicoagellates that have a hollow external skeleton. A ridge about 1.51.7 m wide is central to the main rib which ranged from 2.5 to 4 m wide. The apices and other projections are sometimes decorated with an extensive spiny surface (Fig. 2L) that may merely be a variable feature. It is also possible that these spines continue developing after the basic skeleton is completed (compare Fig. 2F, G). The mechanism of silica deposition in ebriidians is unknown (Preisig, 1994). More is understood about diatom frustules (Schmid & Schultz, 1979) and chrysophycean cysts (Sandgren, 1989) which are formed with the SDV (silica depositional vesicle). Transmission electron microscope studies may elucidate the mechanism of skeleton formation in the asymmetric Hermesinum and determine if a similar membrane is used to make the daughter skeleton. This would require xation of living material. It would also be of interest to determine the timing and mechanism of movement of the newly formed nucleus to the daughter cell. In addition, study of the cellular ultrastructure may help to determine the correct classication of this organism.

Acknowledgements I would like to thank the staff of LFR Levine-Fricke, especially Richard Vogl and Ryan Henry, for the sediment sample; James Watts, Brandon Swan and Kristen Reifel for obtaining water samples; Joan Dainer for technical assistance; and Stuart Hurlbert for reviewing the manuscript. I would like to thank Steven Barlow for the generous use of the Electron Microscope Facility at San Diego State University. I also thank two anonymous reviewers.

References
Bodeanu, N., 1969. Ceretari asupra toplanctonului din zona de mica ad ncime de la litorialul romnese al Mrii Negre. Ecol. Mar. 3: 65147. Carpelan, L. H., 1961. Phytoplankton and plant productivity. In Walker, B. (ed.), The Ecology of the Salton Sea, California in Relation to the Sportshery. Calif. Fish and Game, Fish Bull. 113: 3343. Corliss, J., 1994. An interim utilitarian (user-friendly) hierarchical classication and characterization of the protists. Acta Protozool. 33: 151.

221
Deandre, G., 1952. Classe des bridiens. In Grass, P. (ed.), Trait de Zoologie: Anatomie, Systmatique, Biologie. Masson et Cie, Paris: 407424. Halim, Y., 1960. Observations on the Nile bloom of phytoplankton in the Mediterranean. J. du Cons. 26: 5767. Hargraves, P. & B. Miller, 1974. The ebridian agellate Hermesinum adriaticum Zach. Arch. Protist. 116: 280284. Hasle, G. R. & E. Syvertsen, 1997. Marine diatoms. In Identifying Marine Phytoplankton. Academic Press, Harcourt Brace & Co. San Diego, California. Hovasse, R., 1932. Contribution ltude des silicoagells. Multiplication, variabilit, hrdit, afnits. Bull. biol. Fr. Belg. 66: 447501. Preisig, H., 1994. Siliceous structures and silication in agellated protists. Protoplasma 181: 2942. Rhodes, R. & V. Gibson, 1981. An annual survey of Hermesinum adriaticum and Ebria tripartita, two ebridian algae in the lower Chesapeake Bay. Estuaries 4: 150152. Sandgren, C. D., 1989. SEM investigations of statospore (stomatocyst) development in diverse members of the Chrysophyceae and Synurophyceae. Beih. Nova Hedwigia 95: 4569. Schmid, A. & D. Schultz, 1979. Wall morphogenesis in diatoms: deposition of silica by cytoplasmic vesicles. Protoplasma 100: 267288. Taylor, F., 1990. Incertae Sedis Ebridians. In Margulis, L., J. Corliss, M. Melkonian & D. Chapman (eds), Handbook of Protoctista. Jones & Bartlett Publ., Boston: 720721. Tappan, H. N., 1980. The Paleobiology of Plant Protists. W. H. Freeman & Co., San Francisco: 1028 pp. Vilicic, D., I. Marasovic & G. Kuspilic, 1996/97. The heterotrophic ebridian microagellate Hermesinum adriaticum Zach. in the Adriatic Sea. Arch. Protist. 147: 373379. Zacharias, O., 1906. Eine neue Dictyochide aus dem Mittelmeer, Hermesinum adriaticum n. g., n. sp. Arch. Hydrobiol. und Planktonkunde, Stuttgart 1: 394398.

9152-035-26-28 :xaF 3152-035-26-28 :leT rk.ca.mannohc.mannohc@hokky :rohtua gnidnopserroC*

, , ,  . , . , , . iizluhcs allenisemreH acinecoim airbegnopS uaetalP gnirV , atanoc allenisemreH sutagole sucsinitcA . ijujoH trevlaC . 81 11 5 3 , , 5 2 :
3 snaidirbe ,setallegalfonid lateleksodne ,stsyc ,puorG linoeY TESPXZF, .puorg eht fo noitisoped eht gnirud retaw hsikcarb ro hserf fo ecneulfni eht tseggus stsycotamots sa deredisnoc stsyc naecyposyrhc etunim ,noitidda nI .noitidnoc lanoitisoped ni puorg eht fo eno rewol eht naht redloc tahwemos eb thgim puorg eht fo trap reppu ehT .retaw mraw lanoisacco htiw puorg eht fo noitisoped eht gnirud detanimod sessam retaw dloc taht derrefni si ti ,aera yduts eht fo puorG linoeY eht ni segalbmessa naidirbe dna etallegalfonid lateleksodne ,danomoeahcra gnidulcni tsyc naecyhposyrhc eht morF .puorg eht fo segalbmessa naidirbe eht no desab ,aeS naigewroN ,uaetalP gnirV ni senoz iizluhcs allenisemreH elddiM ot acinecoim airbegnopS fo dna noiger cificaP tsewhtuoS ni senoz atanoc allenisemreH elddiM ot sutagnole sucsinitcA fo slavretni eht htiw detalerroc si puorg eht ,dnA .stsyc danomoeahcra sti yb )napaJ lartneC ni( noitamroF ijujoH eht dna )ASU ni dnalyraM( noitamroF trevlaC eht htiw detalerroc si puorg ehT .ega enecoiM elddiM ot dednopserroc si puorG linoeY eht ,segalbmessa lissoforcim suoecilis evoba no desaB .deifitnedi erew areneg nevele ot gnignoleb seiceps naidirbe neethgie dna areneg eerht ot gnignoleb seiceps etallegalfonid lateleksodne evif ,stsycotamots sa deredisnoc stsyc naecyhpoyrhc fo sepyt eerht rehto dna ,areneg owt ot gnignoleb seiceps danomoeahcra enin ,nisaB gnahoP eht fo trap nrehtron eht ,saera ahgnoehC dna argnoS ni detubirtsid puorG linoeY yraitreT eht morF :tcartsbA

aeroK ,757-005 ujgnawG ,ytisrevinU lanoitaN mannohC ,noitacudE ecneicS fo tnemtrapeD

aeroK ,ecnivorP kubgnoeyK ,nisaB gnahoP eht fo aerA nrehtroN eht ni )yraitreT( puorG linoeY eht morf stsyC lissoF dna puorG lissoforciM suoeciliS roniM
003

*hoK

ooK-gnoeY

,757-005 ,

) 3 (

6002 yraurbeF ,71159 .p ,1 .on ,72 .v ,yteicoS ecneicS htraE naeroK .ruoJ

 auqitna .E . )gninepo( , .)01 ,9 .sgiF ,2 .lP( )bonk( auqitna .E . auqitna sispoirbE )edoirt( 2 . )edalcosem( mutal muidopirtiD )neteleks elbuod( munitores .A . munitores .A . eralugnatcer .A
0 711 911 111 snemiceps latoT 1 2

. munisemreH allenisemreH . iizluhcs .H ,muuqilbo .H ,mucitairda .H munisemreH . atanoc .H allenisemreH . atartsenef allenisemreH munisemreH allenisemreH 4 . . alunirbE )dor evitcennoc( sispoirbE .ps )?(alunirbE . . iessavoh .H munisemreholpaH .)9891 ,initraM dna rekcoL( )telpirt( airbE munisemreholpaH .
63 48 92 573 732 344 512 313 0 3 1 8 9 8 7 2

.)5 .giF ,2 .lP(

1 931 421 58 52 47 2 5 2

4 4 1 1 2 3 2 5 11 61 1 51 1 4 1 9 2 33 1 1

1 2 11 92 1 8 1 4 73 4 7 021 1 1 8 931 3

2 41 4 3 2 72 3 15 1 1 89

1 81 22 32 11

6 2 2 12 11 81 7 72 91 61 4 5 3 63 9 92 4 2 421 36 46 3 4 5 7 591 38 141 2 7

5 2 6 3 4 9 2 3 2 3 2 1 4 31 3 8 2 5 1 1 1 81 11 61 5 7 1 2 13 72 81 4 71 7 2 1 1 3 1 93 55 42 11 03 1 51 5 2 1

2 1 2 3 8 2 2 4 2 1 1 3 1 5 9 7 81 5 31 21 8 2 1 1 51 11 21 1 1 91 12 91 1 6 73 92 33 1 2 1 7 21 6

sepissarc .T .fc muinarhT mutsubor muihcodommaduesP .ps arohpmadoP iregle arohpmadoP sepiunet muinarhtaraP iizluhcs munisemreH muuqilbo munisemreH mucitairda munisemreH atartsenef allenisemreH atanoc allenisemreH iessavoh munisemreholpaH auqitna sispoirbE .ps )?(alunirbE mutal muidopirtiD munitores muihcodommA eralugnatcer .A .fc muihcodommA mucinad muihcodommA allupma muihcodommA
setis gnilpmaS noitamroF axaT

3GI 2GI 1GI 5RJ 4RJ 3RJ 2RJ 1RJ 4DS 3DS 2DS 1DS 3JD 2JD 1JD 1DH 3SH 2SH 1SH noitamroF ohuD noitamroF noejkaH

nisaB gnahoP fo aera nrehtron eht ni puorG linoeY eht morf snaidirbE .2 elbaT

101


31 .giF ,35 .p ,erdnalfeD - .ps .n mucinad muihcodommA 1591 2 .giF ,2 .lP 1591 ,erdnalfeD mucinad muihcodommA

.noitamroF ohuD eht ni rucco ylerar seiceps ehT .drawnwod dehcra syar tsubor sah silibarim .F ,aera yduts eht nI :skrameR .yrtemmys ni laretalib era noteleks eht fo syar ehT .yllacitsiretcarahc ,sesyhpopa lartnec ton sah noteleks ehT .serutcurts raloevla gnivah syar dehcra drawnwod ylhgih evif htiw noteleks depahs rats lanogatneP :noitpircseD 31 ,21 ,4 .sgiF ,2 .lp ,02 ,31 ,21 .sgiF ,1 .lp ,328 .p , cirtimuD - .ps .n silibarim sucsinitcailoF 3791 12 .giF ,1 .lP 3791 , cirtimuD silibarim sucsinitcailoF 3791 , cirtimuD sucsinitcailoF suneG .snoitamrof ohuD eht dna noejkaH eht ni srucco ylerar silicarg .C .tnemegnarra yar sti dna ezis ni elbairav si aera yduts eht ni seiceps ehT .)3791 , cirtimuD( enecoiM elddiM eht ni dnuof saw silicarg .C ,stnemides cificaP nretsewhtuos nI :skrameR .drawnwod dehcra mrof rettel X rednels a fo tsisnoc noteleks eht fo syar ehT .noteleks deyar etatsocirt ruoF :noitpircseD 62 ,12 .sgiF ,4 .lp ,428 .p , cirtimuD - .essavoH silicarg ailofiudraC 3791 01-a9 .sgiF ,94 .lp ,051 .p ,.la te III hcilbeoL - .essavoH silicarg ailofiudraC 8691 01 .giF ,721 .p ,essavoH - .ps .n silicarg ailofiudraC a2391 91 .giF ,1 .lP 2391 ,essavoH silicarg ailofiudraC 2391 ,essavoH ailofiudraC suneG

.snoitamrof ohuD eht dna noejkaH eht ni srucco seiceps ehT .)5791( nesleiN -hcreP yb snemiceps sti fo esoht naht swodniw rediw tahwemos evah puorG linoeY eht morf allupma .A fo snemiceps ehT .)5791 ,nesleiN-hcreP( cificaP nretsewhtuoS morf seroc fo slavretni enecoE ot enecogilO ni derrucco osla tI .)4391 ,erdnalfeD( dnalaeZ weN fo stisoped enecoE reppU eht morf detroper yllaitini saw allupma .A :skrameR .ylevitcepser ,gnir lacipatna ro lacipa dna meht neewteb snoitcnuj ta swodniw rewol dna reppu fo tsisnoc sedalcohtsipo dna sedalcorp ehT .senitca morf dehcnarb era sedalcohtsipo dna sedalcorp ,noteleks eht nI .senitca eerht dna sedalcohtsipo eerht ,sedalcorp eerht ,gnir lacipatna na ,gnir lacipa na fo gnitsisnoc noteleks edoirt ralugnatceR :noitpircseD 42 ,32 .sgiF ,1p .lp ,reniatruoF dna oppilifnaS - .erdnalfeD allupma muihcodommA 3002 62-32 .sgiF ,5 .lp ,91-71 ,sgiF ,4 .lp ,nesleiN -hcreP - .erdnalfeD allupma muihcodommA 5791 1 .giF ,84 .lp ,41 .giF ,.la te III hcilbeoL - .erdnalfeD allupma muihcodommA 8691 911 .giF ,821 .p - .erdnalfeD allupma muihcodommA 2591 2 .giF ,77 .p ,erdnalfeD - .ps .n allupma muihcodommA 4391 1 .giF ,2 .lP 4391 ,erdnalfeD allupma muihcodommA 2391 ,essavoH muihcodommA suneG 0591 ,erdnalfeD eaecaihcodommA ylimaF 4691 ,.la te grebinroH SELAIRBE redrO 0791 ,III hcilbeoL EAECYHPOIRBE ssalC 4191 ,rehcsaP ATYHPOHRRYP noisiviD
SNAIDIRBE

.snoitamrof ohuD eht dna noejkaH eht ni erar yrev si seiceps ehT .)3791( cirtimuD yb sairetsartet .A fo esoht ot elbarapmoc ,noitcerid dehcterts ni ralugerri tahwemos dna dehcra ylhgih era aera yduts eht ni sairetsartet .A fo syar ehT :skrameR .serutcurts raloevla kaew derevoc ro htooms si syar eht fo ecafruS .etalp ekil dimaryp ralugnatcer a htiw noteleks deyar ruoF :noitpircseD

901


.5RJ ,noitamroF ohuD ,004 .initraM dna rekcoL )erdnalfeD( atanoc allenisemreH .21 .giF .2RJ ,noitamroF ohuD ,004 .initraM dna rekcoL iessavoh munisemreholpaH .11 .giF .2SH ,noitamroF ohuD ,acirol detaluspacne na htiw nemiceps ,004 .essavoH )zluhcS( auqitna sispoirbE .01 .giF .3SH ,noitamroF noejkaH ,nemiceps deificilisrepyh ,004 .essavoH )zluhcS( auqitna sispoirbE .9 .giF .2DS ,noitamroF ohuD ,004 .essavoH )zluhcS( auqitna sispoirbE .8 .giF .2DS ,noitamroF ohuD ,004 .ps )?(alunirbE .7 .giF .3SH ,noitamroF noejkaH ,004 .essavoH mutal muidopirtiD .6 .giF .2DS ,noitamroF ohuD ,noteleks elbuod ,004 .initraM dna rekcoL munitores muihcodommA .5 .giF .1RJ ,noitamroF ohuD ,004 .initraM dna rekcoL munitores muihcodommA .4 .giF .1SH ,noitamroF noejkaH ,004 .erdnalfeD )zluhcS( eralugnatcer .A .fc muihcodommA .3 .giF .2DS ,noitamroF ohuD ,006 .erdnalfeD mucinad muihcodommA .2 .giF .3RJ ,noitamroF ohuD ,004 .erdnalfeD allupma muihcodommA .1 .giF snaidirbE

2 etalP

011

5-4 .sgiF ,2 .lP 6891 ,initraM dna rekcoL munitores muihcodommA .snoitamrof ohuD dna noejkaH eht ni erar yrev era eralugnatcer .A .fc muihcodommA fo snemiceps ehT .)6891( initraM dna rekcoL yb debircsed eralugnatcer .A ot ralimis swodniw roiretna ralugnatcer dna edalcnys depahs T a evah yeht ,dnA .sedalcohtsipo dna sedalcorp evissam gnivah ,munitores .A morf dehsiugnitsid era yeht tuB .eralugnatcer muihcodommA elbmeser ylesolc snemiceps naidirbe ehT :skrameR 3 giF ,2 .lP 2391 ,)zluhcS( eralugnatcer .A .fc muihcodommA .snoitamrof ohuD eht dna noejkaH eht ni erar yrev si seiceps ehT .yduts siht ni mucinad .A sa deifissalc saw noxat siht ,eroferehT .)5791( nesleiN-hcreP yb debircsed mutruc mucinad .A morf tnereffid era aera yduts eht fo snemiceps eht ,sedalc xevnoc drawtuo gnivah ,tuB .)5791( nesleiN -hcreP yb detroper mutruc mucinad .A ot ralimis era yduts eht morf snemiceps mucinad .A :skrameR .poleved ton od seno rewol tub trap lacipa ni detacol era swodniw reppu ,noteleks eht nI .drawni devruc gnir lacipatna na htiw noteleks edoirt cibuc dednuor llamS :noitpircseD 4-3 ,sgiF ,5 .lp ,088 .p ,nesleiN-hcreP - .erdnalfeD mutruc mucinad muihcodommA 5791 c9-2 .sgiF ,84 .lp ,441 .p ,.la te III hcilbeoL - .erdnalfeD mucinad muihcodommA 8691

.oot ,sedalcosem nekorb wohs aera yduts eht morf mutal .D fo snemiceps tsoM .ylisae ffo nekorb eb ot dnet mutal muidopirtiD fo sedalcosem eht ,)6891( initraM dna rekcoL ot gnidroccA :skrameR .sedalcosem gnidulcni sgninepo owt dna ,sedalcotsipo trohs detacrufib ,gnir lacipa egral htiw noteleks epyt etteugaB :noitpircseD 6 ,5 .sgiF ,2 .lp ,349 .p ,initraM dna rekcoL - .essavoH mutal muidopirtiD 6891 01 .giF ,73 .lp ,351 .p ,.la te III hcilbeoL - .essavoH mutal muidopirtiD 8691 6 .giF ,282 .p ,essavoH - .ps .n mutal muidopirtiD b2391 6 .giF ,2 .lP 2391 ,essavoH mutal muidopirtiD 2391 ,essavoH muidopirtiD suneG 1591 ,erdnalfeD eaecaidopirtiD ylimaF .snoitamrof ohuD eht dna noejkaH eht ni tnadnuba yrev era seiceps ehT .)6891 ,initraM dna rekcoL( eralugnatcer .A ot elbarapmoc ,sgninepo denoitnem evoba eht dna ,sedalcnys dehcra ylhgih eht ,noteleks elicarg erom sti yb dehsiugnitsid si munitores .A :skrameR .sedalc gnidnuorrus dna edoirt a neewteb sgninepo roiretna ralucricimes sah noteleks ehT .sedalcohtsipo dna sedalcorp fo spit eht ni swodniw ralugnairt ,sedalcnys devruc ylgnorts htiw noteleks edoirt ralugnatceR :noitpircseD 2 ,1 .sgiF ,2 .lp ,349 .p ,initraM dna rekcoL - .ps .n munitores muihcodommA 6891

.3SH ,noitamroF noejkaH ,004 .essavoH sepissarc .T .fc muinarhT .22 .giF .1SH ,noitamroF noejkaH ,004 .erdnalfeD mutsubor muihcodommaduesP .12 .giF .2SH ,noitamroF noejkaH ,acirol detaluspacne na htiw nemiceps ,004 .tdrahniemeG iregle arohpmadoP .02 .giF .2SH ,noitamroF noejkaH ,004 .ps arohpmadoP .91 .giF .2DS ,noitamroF ohuD ,noteleks elbuod ,004 .essavoH sepiunet muinarhtaraP .81 .giF .2DS ,noitamroF ohuD ,004 .essavoH sepiunet muinarhtaraP .71 .giF .3RJ ,noitamroF ohuD ,004 .essavoH iizluhcs munisemreH .61 .giF .1RJ ,noitamroF ohuD ,006 .initraM dna rekcoL muuqilbo muisemreH .51 .giF .2DS ,noitamroF ohuD ,006 .sairahcaZ mucitairda munisemreH .41 .giF .2RJ ,noitamroF ohuD ,004 .illeugnerF atartsenef allenisemreH .31 .giF .deunitnoC .2 etalP

111


.elicarg yllareneg era noteleks eht fo sedalc ehT .eerged ytxis tuoba fo etats dessorc eht ni denioj yllautum era sdopirt owt ehT .snoitatnedni emos htiw sedalc devruc drawni yb detcennoc sdopirt owt fo desopmoc noteleks depahs snel ralucriC :noitpircseD 51-41 .sgiF ,2 .lp ,449 ,349 .p ,initraM dna rekcoL - .)gniL( atunroc sispoirbE 6891 51 .giF ,4 .lp ,088 .p ,nesleiN-hcreP - .)zluhcS( auqitna sispoirbE 5791 81 ,71 .sgiF ,3 .lp ,512 .p ,gniL - .)zluhcS( auqitna auqitna sispoirbE 7791 1 .giF ,021 .p ,essavoH - .bmoc .von )zluhcS( auqitna sispoirbE a2391 f-a96 .sgiF ,372 .p - .zluhcS auqitna airbE 8291 01-8 .sgiF ,2 .lP 2391 ,)zluhcS( auqitna sispoirbE 2391 ,essavoH sispoirbE suneG 3491 ,essavoH eaecanisemreH ylimaF .snoitamrof ohuD eht dna noejkaH eht ni erar era yehT .eno roiretsop htiw edalcnys roiretna gnitcennoc dor evitcennoc fo ecnetsixe eht ni axodarap .E ot ralimis era aera yduts eht morf mrof tneserp eht fo snemiceps ehT .)0591 ,erdnalfeD( uramaO fo stisoped enecoE eht morf detroper yllaitini saw ,alunirbE suneg fo seiceps euqinu a ,axodarap .E :skrameR 7 .giF ,2 .lP .ps )?(alunirbE 0591 ,erdnalfeD alunirbE suneG 1091 ,nnamremmeL eaecairbE ylimaF .noitamroF noejkaH eht ni ylno srucco mrof ehT .snroh drawni llams gnivah sedalcohtsipo dehcterts ylthgiarts yb sepissarc .T morf dehsiugnitsid si mrof eht tuB .eniltuo ni sepissarc muinarhT ot ralimis si mrof sihT :skrameR 22 .giF ,2 .lP 2391 ,essavoH sepissarc .T .fc muinarhT 2391 ,essavoH muinarhT suneG

.snoitamrof ohuD eht dna noejkaH eht ni erar si seiceps ehT .mutsubor .P lacipyt fo esoht ot evitaler ,regral ssel ro erom era serop laicifrepus ,aera yduts eht morf mutsubor muihcodommaduesP fo nemiceps eht roF :skrameR .noteleks eht fo pot eht ta decalp dna llams si gninepo lacipa ehT .gninepo lacipa na dna ecafrus detarofrep a htiw noteleks diospillE :noitpircseD 8-6 .sgiF ,6 .lp ,375 .p ,initraM dna rekcoL - .erdnalfeD mutsubor muihcodommaduesP 9891 b81-61 .sgiF ,34 .lp ,581 ,481 .p ,.la te III hcilbeoL - .erdnalfeD mutsubor muihcodommaduesP 8691 24-93 .sgiF ,49 .p ,erdnalfeD- .ps .n mutsubor muihcodommaduesP 4391 12 .giF ,2 .lP 4391 ,erdnalfeD mutsubor muihcodommaduesP 2391 ,essavoH muihcodommaduesP suneG .snoitamrof eht ni deretnuocne osla era smrof noteleks elbuod sti ,dnA .snoitamrof ohuD eht dna noejkaH eht ni erar si seiceps ehT .sedalcohtsipo devruc drawtuo ylthgils dna wodniw reppu xevnoc-drawpu na evah snemiceps eht esuaceb )9891 ,6891( initraM dna rekcoL yb desoporp mutarhtalc .P morf tnereffid era aera yduts eht morf sepiunet .P fo snemiceps ehT :skrameR .drawtuo devruc yllanoisacco era noteleks eht fo sedalcohtsipo ehT .sedalcohtsipo dehcterts drawnwod gnol dna gnir lacipa na htiw noteleks depahs lootS :noitpircseD 71-61 .sgiF ,74 .lp ,971 .p ,.la te III hcilbeoL - .essavoH sepiunet muinarhtaraP 8691 564 ,464 .p ,essavoH - .bmoc .von sepiunet muinarhtaraP c2391 5 .giF ,321 .p - .essavoH sepiunet minarhT a2391 81 ,71 .sgiF ,2 .lP 2391 ,essavoH sepiunet muinarhtaraP 2391 ,essavoH muinrhtaraP suneG .snoitamrof ohuD eht dna noejkaH eht ni srucco ylerar seiceps ehT

211

ni detacol si wodniw llams a ,noteleks eht fo trap roiretna eht nI .strap lacipatna dna lacipa ni senips owt htiw noteleks eneairt epyt etiK :noitpircseD 2-1 .sgiF ,022-812 .p ,ynaffiT - .sairahcaZ mucitairda munisemreH 2002 6-5 .sgiF ,1 .lp ,449 .p ,initraM dna rekcoL - .sairahcaZ mucitairda munisemreH 6891 01-a9 .sgiF ,04 .lp ,961-861 .p ,.la te III hcilbeoL - .sairahcaZ mucitairda munisemreH 8691 d-a .sgiF ,493 .p ,sairahcaZ - .ps .n mucitairda munisemreH 6091 41 .giF ,2 .lP 6091 ,sairahcaZ mucitairda munisemreH 6091 ,sairahcaZ munisemreH suneG

eht morf atanoc .H fo nemiceps ehT :skrameR .htgnel ni tnereffid yllautum era noteleks eht fo sedalcohtsipo ehT .noteleks eht fo etalp roiretna eht ni detacol si wodniw reppu llams A .sedalcnys devruc ylgnorts htiw noteleks eneairt dednuor llamS :noitpircseD 01-9 .sgiF ,2 .lp ,449 .p ,initraM dna rekcoL .bmoc .von )erdnalfeD( atanoc allenisemreH 6891 141 .giF ,86 ,64 ,44 .p - .erdnalfeD mutanoc munisemreH 1591 21 .giF ,2 .lP 6891 ,)erdnalfeD( atanoc allenisemreH 4391 ,erdnalfeD allenisemreH suneG .snoitamrof ohuD eht dna noejkaH eht ni srucco ylerar seiceps ehT .sedalcnys thgiarts retrohs dna evissam erom swohs aera yduts eht ni eno eht ,)9891( initraM dna rekcoL yb iessavoh .H fo nemiceps eht htiw gnirapmoC :skrameR .dnuor si eno rehto dna thgiarts era owt trohs ,noteleks eht fo sedalcnys eht fO .edalcohtsipo na dna ,edalcorp a ,sedalcnys owt fo desopmoc telpirt demrof Y citsiretcarahc a htiw noteleks depahs snel ralucriC :noitpircseD 21-11 .sgiF ,7 .lp ,175 .p ,initraM dna rekcoL - .ps .n iessavoh munisemreholpaH ,9891 11 .giF ,2 .lP 9891 ,initraM dna rekcoL iessavoh munisemreholpaH 3491 ,essavoH munisemreholpaH suneG

.snoitamrof ohuD eht dna noejkaH eht ni srucco ylerar seiceps ehT .)6891 ,initraM dna rekcoL( cificaP nretsewhtuos ni enecoiM reppU eht fo trap rewol eht ot enecoiM elddiM eht ni detubirtsid si atartsenef .H .)6891( initraM dna rekcoL fo atanoc allenisemreH fo taht ot elbarapmoc ,sedalc dna edbahr a neewteb noitcnuj htooms dna eniltuo ni ralucric ylraen si aera yduts eht ni nemiceps ehT :skrameR .ertnec sti ni wodniw ralucric llams a htiw etalp ralugnairt a ekam noteleks eht fo trap roiretna eht ni sedalC .dehcra ylhgih sedalc esogur dna tsubor htiw noteleks eneairt ralucric llamS :noitpircseD 31 .giF ,2 .lp ,449 .p ,initraM dna rekcoL - .illeugnerF atartsenef allenisemreH 6891 9 .giF ,15 .lp ,461 .p ,.la te III hcilbeoL - .illeugnerF atartsenef allenisemreH 8691 a5 .giF ,972 .p ,illeugnerF - .ps .n atartsenef allenisemreH 1591 31 .giF ,2 .lP 1591 ,illeugnerF atartsenef allenisemreH .snoitamrof ohuD dna noejkaH eht ni srucco yltneuqerf atanoc .H .eniltuo lateleks dna tnemegnarra naidalcohtsipo ni elbairav tahwemos era aera yduts eht ni snemiceps ehT .ylevitarapmoc ,)6891( initraM dna rekcoL fo taht htiw dedicnioc llew era aera yduts

.ecnerrucco ni snoitamrof ohuD eht dna noejkaH eht ni tnadnuba dna tnetsisnoc si seiceps ehT .auqitna .E ot dengila erew sedalc detnedni ylralugerri htiw snemiceps ,yduts siht nI .mir delknirw niht a yb dednuorrus si gninepo ehT .gninepo ediw a htiw epahs derraj derepat yltneg a swohs smrof etacirol eht ni aciroL .smrof gniraeb acirol dna ,snoteleks rieht ni xepatna ro/dna xepa eht ta senips llams htiw snemiceps ,smrof deificilisrepyh ,sedalc depoleved ylkciht htiw snemiceps sa seiteirav lareves swohs auqitna .E ,aera yduts eht nI :skrameR

311


ylbaborp

si

gnir

lacipA

.elicarg .P
a sah

ot

elbarapmoc dna teksab eht tuB aera

tcnitsid

ekam

enitca

na

morf

detacrufirt

sedalc

,snoitatnedni naidalcohtsipo .ygolohprom yduts eht

tuohtiw ediw

sedalc

htooms

,noteleks eht fo trap roiretsop eht nI .sedalc devruc ylgnorts htiw noteleks eneairt diovO :noitpircseD

rehtar

nemiceps

ni

elicarg arohpmadoP
nemiceps naidirbe 91 .giF ,2 .lP .ps

selbmeser

)41(3 .giF .181 .p ,yentraCcM dna eessinrE - .essavoH

morf

sihT

:skrameR

iizluhcs munisemreH

5991

02-01 .sgiF ,14 .lp ,471 .p ,.la te

arohpmadoP
si seiceps ehT .tsyc

III

hcilbeoL - .essavoH

8691 a2391

521 .p ,essavoH - .ps .n

iizluhcs munisemreH iizluhcs munisemreH iizluhcs munisemreH

swohs tnereffid

.ecnerrucco ni noitamroF noejkaH eht fo trap rewol eht ni detcirtser

61 .giF ,2 .lP 2391 ,essavoH

ton eb dluow snaidirbe ni eacirol eht taht detseggus )0891( nappaT tuB .)5991 ,yentraCcM dna eessinrE( sserts a ni latnemnorivne yam tI fo gnirud eht elor sah evitcetorp osla .)1391 a tsom yb nI aera fo dnik eht eht ni .snoitamrof ohuD eht dna noejkaH secnerrucco ni tnetsisnoc lacipa

muuqilbo
yllautum rekcoL( talf

evres

.acirol

yduts

.H

.htgnel

senips

iregle .P
sti

nemiceps

ehT raj

,tdrahniemeG( deziretcarahc :skrameR

owt ro sedalcnys devruc yllanoisacco sah aera yduts eht ni seiceps ehT .)6891 ,initraM dna

xepa si

ta

acirol

depahs

iregle arohpmadoP
sedalc raj a

,sesac

enips

lacipa

lacirtemmysa

dna

sedalcnys

reppu

.snoitaluciter ralugerri yb derevoc si kcen trohs a htiw noteleks eht fo acirol ehT .devruc ylhgih depahs detnedni htiw dna xepa eneairT eht ta acirol

yllanigiro

sedulcni

muuqilbo munisemreH
gnir lacipa no

:skrameR

.swodniw fo tsisnoc ton od noteleks eht fo sedalcohtsipo ehT .drawnwod detlit sedalcohtsipo derepat owt dna enips gnol

noteleks

:noitpircseD

)51(3 .giF ,181 .p ,yentraCcM dna eessinrE - .tdrahniemeG

lacirtemmysa a htiw noteleks eneairT :noitpircseD 4 ,1 .sgiF ,1 .lp ,449 .p ,initraM dna rekcoL - .ps .n

iregle arohpmadoP

5991

71-61 .sgiF ,7 .lp ,275 .p ,initraM dna rekcoL - .tdrahniemeG

muuqilbo munisemreH

6891

iregle arohpmadoP

9891 51 .giF ,2 .lP 6891 ,initraM dna rekcoL 8691

4 .giF ,44 .lp ,281 .p ,.la te III hcilbeoL - .tdrahniemeG

iregle arohpmadoP

muuqilbo munisemreH
.yduts siht

91 .giF ,01 .lp ,701 .snoitamrof ohuD eht dna noejkaH 1391 eht ni srucco yltnetsisnoc

.p

,tdrahniemeG - .ps

.n

iregle arohpmadoP

mucitairda .H
sti

02 .giF ,2 .lP 1391 ,tdrahniemeG 1391 ,tdrahniemeG

ni dnuof semitemos era seiceps eht fo segats eavral sa deredisnoc seneairt rellamS .decuder dna dedorroc yltneuqerf ylevitaler era si sedalcohtsipo enips a lasab sti ,dnA .tcnitsidni lacipa

iregle arohpmadoP arohpmadoP suneG

.snoitamrof ohuD yltnetsisnoc roiretna dehcra ni seiceps wodniw dna

tuB

.enips eht

gnol nI

ylbakramer

sedulcni dna

seiceps ,6891( ,sedalc

,noitidda dna

eht ehT

dna

noejkaH

eht

ni si

srucco trap

.)2002( fo

ynaffiT taht ot

)9891

initraM dna eht

.diovo egral

tahwemos stI

rekcoL a ni

elbarapmoc si

edbahr morf

reppu

.sedalcohtsipo htiw

drawtuo a

htooms

ylnommoc

aera

yduts

snoitatnedni aera yduts

suoivbo eht morf

telpirt

tcnitsid ehT

swohs

mucitairda .H
fo eno eht si

ehT .)3991 ,yentraCcM dna eessinrE(

iizluhcs .H
Y

:skrameR

tneserp eht litnu devivrus seiceps naidirbe erar yrev

.edbahr ot noitcennoc dehcra ylhgih dna mrof rettel swohs noteleks eht

mucitairda munisemreH

:skrameR .sedalc

fo telpirt roiretsop ehT .wodniw llams a htiw telpirt

fo detsisnoc erutcurts demrof T eht fo ertnec eht

411

-ruoJ .aeroK ,nisaB gnahoP eht fo trap lartnec eht morf stsyc etallegalfonid enecoiM ,2991 ,.H ,nuY dna .H ,nuyB .p 912 ,ytisrevinU lanoitaN luoeS ,sisehT .D .hP .nisaB yratnemideS gnahoP eht ni atartS eneg -oeN eht fo yhpargitartS dna ygolonylaP ,5891 ,.Y .P ,gnoB .451-731 ,)2(31 ,aeroK fo yteicoS lacigol -otnoelaP eht fo lanruoJ .aeroK ,nisaB gnahoP nrehtuos eht ni )enecoiM elddiM( noitamroF noejgaH eht morf sanuaf nairaloidaR ,7991 ,.H ,nuY dna .D .J ,eeL ,.Y ,kaB .162-522 ,)2(21 ,aeroK fo yteicoS lacigolotnoelaP eht fo lanruoJ .aeroK ,nisaB gnahoP eht ni noitamroF ohuD eht morf snairaloidaR enecoiM elddiM ,6991 ,.H ,nuY dna .D .J ,eeL ,.Y ,kaB .811-78 ,9 ,ytisrevinU awazanaK fo tropeR ecneicS ,napaJ lartneC ,alusnineP otoN ,enots -dum suoecamotaid ijujoH enecoiM eht ni sdanomo -eahcra dna setallegalfocilis ,serops dna sniarg nellop ,smotaid lissoF ,.la te ,.W ,awakihcI nI .eaecadanomo -eahcrA dnu edillegalfociliS II traP ,4691 ,.A ,nnamhcaB .448-938 ,29 ,I traP ,nitelluB aciremA fo yte -icoS lacigoloeG .srotacidni latnemnorivne laitnetop sa stsyc etyhposyrhC ,1891 ,.D .A ,doohaM dna .P .D ,madA

.122-512 ,)3(61 , . )( ,5991 , , , .26-23 ,)1(7 , . ,1991 , , , .09-46 ,)1(6 , . 3 ,0991 , .192-382 ,83 , . 3 ,4891 , , .p 208 , . ,9991 , , , , , , , , , , , , , , , , .p . ,0002 , , 022 ,

, .

4002

.noitamroF noejkaH eht fo trap rewol eht ylno ni rucco yehT .nemiceps eht ni decuder

led soiraloidaR y sodalegalfocilS ,1491 ,.J ,illeugnerF .211-73 ,7 ,aigolotnoelaP ,2 ,2 eireS aveuN ,atalP aL ed oesuM led atsiveR .selisf sod -alegalfocils sol erbos senoicaredisnoC ,0491 ,.J ,illeugnerF .581-771 ,8 ,ygolotnoelaP ni sesruoC trohS yte -icoS lacigolotnoelaP ,slissoforcim suoeciliS ,)sronevnoc( .la te .D .C ,emolB nI .setallegalfonid lateleksodne dna snaidirbE ,5991 ,.K ,yentraCcM dna .J .J ,eessinrE .041-131 ,snoitacilbuP cifitneicS llewkcalB .stsitorp dna setoyrakorp lissoF ,).de( .H .J ,sppiL nI ,snaidirbE ,3991 ,.K ,yentraCcM dna .J .J ,eessinrE .001-88 ,)3(02 ,setoN cigoloeG .aniloraC htuoS ,ytnuoC repsaJ ,yalC eihctahwasoC eht morf setallegalfonid lateleksodnE ,6791 ,.J .J ,eessinrE .p 473 ,ssoV dlopoeL ,gizpieL ,edrE red fua snebeL negidnts -bles nenielk rabthcisnU sed nekriW edneffahcs nesleF dnu nedrE sad .eigoloegorkiM ,4581 ,.G .C ,grebnerhE .p 981 .thcerd -roD ,srehsilbuP cimedacA rewulK .stsyC naecyhp -osyrhC fo saltA ,5991 ,.P .J ,lomS dna .B ,beeZ ,.K ,ffuD .538-918 ,12 ,PDSD fo stropeR laitinI .PDSD eht fo 12 geL gnirud deroc stnemides cificaP nretsewhtuoS eht ni setallegalfonid lateleksodne ciozoneC ,3791 ,.P , cirtimuD .142-722 ,)1(31 ,eigoloeG aireS ,eifargoeg , cizifoeg ,eigoloeg ed iratecrec i iidutS .ecitaprac iinuiger lunainotroT nid ir loidar uc soligra iulutnoziro arup -usa ecigolotnoelaporcim iitaredisnoC ,8691 ,.P , cirtimuD .821 -521 ,1 ,eiC te nossaM ,siraP ,eigolotnoelaP ed tiarT ,.J ,uaeteviP ni .snidirb sed essalC ,2591 ,.G ,erdnalfeD .48-1 ,58 ,euqigleB al ed te ecnarF al ed euqigoloiB nitelluB .euqitamtsyS ,noitulov ,eigolo -ibolaP ,snidirb sel rus sehcrehceR ,1591 ,.G ,erdnalfeD .2871-0871 ,032 ,secneicS sed eimdacA'l ed sudneR setpmoC .eadihcodommA el ceva erneg ec ed snoit -aler te airbE'd etteleuqs ud esylanA ,0591 ,.G ,erdnalfeD .p 89 ,533 ,selle -irtsudnI te seuqifitneicS stilautcA ,eiC & nnamreH ,siraP ,euqigolog eloR ,euqigolotnolap te euqigoloib urepA .selissof sllegalf seL ,6391 .G ,erdnalfeD .69-57 ,4 ,eigolotsitorP ed selannA .sellevuon semrof seuqleuq ed noitpircsed te seca -irb sed etteleuqs ud erutalcnemoN ,4391 .G ,erdnalfeD .553-643 ,)6-5(67 ,ecnarF ed euqinatoB ticoS al ed nitelluB .sec -danomoeahcrA sel rus eton ednoceS ,3391 ,.G ,erdnalfeD .122-512 ,)2(02 ,aeroK fo yteicoS lacigoloeG eht fo lanruoJ ehT .aeroK fo snoitamrof )enecoiM( gnahoP dna gnodI morf sdi -tinamsaT ,4891 ,.K .B ,miK dna .Y .P ,gnoB ,.K .D ,iohC .532 -461 ,)2(8 ,aeroK fo yteicoS lacigolotnoelaP eht fo lan

511


Middle Eocene ebridians from the central Arctic Basin

2

Center for Advanced Marine Core Research, Kochi University, B200 Monobe, Nankoku, 783-8502, Japan Department of Earth and Planetary Sciences, Kyushu University, Hakozaki6-10-1, Fukuoka, 812-8581, Japan email: jm-jo@kochi-u.ac.jp; kozo@geo.kyush-u.ac.jp

Jonaotaro Onodera1 and Kozo Takahashi2

ABSTRACT: Abundant and diversified ebridians recovered during IODP Expedition 302 (ACEX) have been identified and counted in order to establish their taxonomy and to decipher the biostratigraphic potential of ebridians in the central Arctic Ocean. In the ACEX samples these fossils are preserved in Lithologic Units 1/6 and 2, which consist mainly of dark silty clay and biosiliceous ooze, respectively. Thirty taxa have been distinguished, three of which are described as new species (Ammmodochium lomonosovense, Pseudammodochium karyon, and Pseudammodochium psichion). The most dominant ebridian species is Pseudammodochium dictyoides throughout the biosiliceous section. The second dominant species varies alternately throughout the section. Based on the characteristic occurrences of major ebridian taxa, the ebridian assemblages were divided into Groups A to D in stratigraphic order. The ebridian assemblages in piston core USGS Fl-422 from the Alpha Ridge probably correlate to our assemblage Group A of early middle Eocene age, although rare younger taxa are irregularly included.

INTRODUCTION

Ebridians represent a group of marine zooflagellates that have two dissimilar flagella and a siliceous internal skeleton. The ebridians are cosmopolitan but usually rare in most marine plankton assemblages, with several exceptions in specific conditions such as in estuaries, bays, and semi-closed seas such as the Baltic, and the Black Sea (Tappan 1980, Earnissee and McCartney 1993, Korhola and Smol 2001, Hargraves 2002, Osawa et al. 2005). Previous studies of Eocene sediments in the Arctic Ocean were conducted only based on piston core samples. Thus the paleoceanographic and paleontological understanding of the Eocene Arctic Ocean remained fragmentary (Clark 1974, Kitchell and Clark 1982, Bukry 1984, Ling 1972, 1985a, Clark et al. 1986, Sims and Ross 1988, Dellagnese and Clark 1994, Magavern et al. 1996). The IODP Expedition 302 (Arctic Coring Expedition: ACEX) successfully cored middle Eocene sediments in the central Arctic Ocean in the summer of 2004 (Backman et al. 2006, 2008, Moran et al. 2006). This successful continuous coring made high resolution studies of the Eocene Arctic Ocean sediments possible. The previous studies and the recent ACEX investigations demonstrated that the Eocene sediments from the Arctic Ocean contain siliceous microfossils assemblages mainly consisting of diatoms, silicoflagellates, chrysophyte cysts and ebridians (Bukry 1984, Ling 1985a, Dellagnese and Clark 1994, Stickley et al. 2008). Radiolarians are nearly absent in the ACEX samples (Backman et al. 2006). Ebridians represent one of the significant groups of siliceous microfossil assemblages in the Eocene Arctic Ocean (Dellagnese and Clark 1994, Backman et al. 2006, Stickley et al. 2008). The age determination potential of ebridians was generally limited to the level of epoch for the Paleogene (Loeblich et al. 1968), because the standard ebridian biostratigraphy has only been established for the Neogene (Locker and Martini 1986; Ernissee and McCartney 1993) except for a single study of the Eocene and Oligocene in the Southern Ocean (Bohaty and Harwood 2000). Consequently, the highly abundant and diversified occurrences

of ebridians in the ACEX samples are very important. It was expected that the continuous Eocene sediment core samples from the Arctic Ocean obtained by the IODP Expedition 302 would be able to improve the ebridian taxonomy and biostratigraphy. For this reason, this paper focuses on the ebridian taxonomy in the ACEX cores.
MATERIALS AND METHODS

Siliceous sediments were retrieved from Sites M0002 and M0004 on the Lomonosov Ridge in the central Arctic Ocean by the ACEX IODP Expedition 302 in summer 2004 (text-fig. 1; Table 1). In the Lithologic Units of the ACEX cores, the studied samples from Hole M2A represent Lithologic Unit 1/6 and the upper part of Unit 2, whereas the samples from Hole M4Arepresent mainly the lower part of the Lithologic Unit 2 (text-fig. 2). By combining the available samples from both sites it is possible to study the Eocene siliceous fossiliferous section as one continuous sequence. The samples and methods for this study are the same as those for the silicoflagellate study by Onodera and Takahashi (in this volume) and are described below. Sample preparation and counts A total of 122 core-sediment and 29 core catcher samples from Holes M2A and M4A were quantitatively processed in order to determine the abundances of ebridian species. For biostratigraphy and paleoceanographic reconstruction, samples were usually taken at 20cm intervals in each section. The samples were freeze dried, measured for dry weight, and then treated with hydrogen peroxide, hydrochloric acid, and Calgon (mainly composed of sodium hexametaphosphate). In the chemical treatment process, ultra sonic was applied for 10 seconds in order to disaggregate consolidated clay particles. After adequately diluting the acid and other chemicals by decantation several times, the remaining sample material was sieved through a 45m mesh, and then both fractions were filtered through 0.45m nominal pore size membrane filters. The filtered volume of the fine fraction is 1/20 of total volume of the material. Dried membrane filters were mounted with Canada balsam onto glass slides.
187

micropaleontology, vol. 55, nos. 2-3, pp. 187-208, text-figures 1-2, plates 1-7, tables 1-2, appendix 1, 2009

Micropaleontology, vol. 55, nos. 2-3, 2009

TABLE 2

Mean abundance of ebridian taxa in each ebridian assemblage group in this study. The symbol + represents the mean abundance of <1% in total ebridians.

TAXONOMY

The ebridian species in the ACEX samples belong to 26 taxa. These taxa are described or discussed in alphabetical order. Their morphological terminologies are mainly based on Deflandre (1934). Class Ebriophyceae Loeblich III, 1970 Order EBRIALES, Honigberg et al. 1964 Genus Ammodochium Hovasse 1932c
Ammodochium complexum Dumitrica and Perch-Nielsen 1976

Derivation of Name: The specific name lomonosovense is derived from the Lomonosov Ridge of the central Arctic Ocean, where IODP Expedition 302 drilled and cored the samples which contain the present taxon. Holotype: Plate 1, figure 11a, b, Sample IODP302-M2A61X-2, 76-78cm, slide MPC-02687 (Micropaleontology Collection, National Science Museum, Tokyo), England finder K21/3. Type locality: IODP302 Site M0002 Hole A, 264.76-264.78 mbsf, Lomonosov Ridge, central Arctic Ocean. Description: Skeleton ovoid in dorsal-ventral view with two rings of different diameters. Each bifurcated proclade and opisthoclade with three large pores of which one, in both the proclade or opisthoclade, is located near the distal end of the actine, and two are collaterally located on the ring, at the distal end of the clades. Anterior and posterior synclade very broad with three pairs of pores, each pair located at the connection between the proclades and opisthoclades and the anterior and posterior synclades, respectively. In highly silicified species the posterior synclades are so broad that their inner borders reach the core of the triactine. In this case in lateral view, the SEM and LM images (Pl. 2, Figs. 2 and 3) show a wide central pore (between the triactine spicule and the anterior synclade) bordered by eight pores. Of these pores two pairs are on the synclades and two pairs on the proclades and opithiclades respectively. Dimensions: Length 20-33m; width 18-33m [N = 25]. Occurrence: From Samples M4A-15X-CC, 24-26cm to M2A53X-1, 36-38cm. This taxon usually co-occurs with A. fletcheri and the silicoflagellate Dictyocha arctios. Remarks: This species is similar to the holotype of Ammodochium complexum, from which it differs in having no pore between the mesoclades and the ends of actins and in having a heavily silicified skeleton. It differs from A. novum Perch-Nielsen by the absence of the extra pore between the proclade and opisthoclade.
191

Plate 1, figure 1
Ammodochium complexum Dumitrica & Perch-Nielsen, in PERCHNIELSEN, 1976, p. 152, pl. 8, figs. 15, 16; pl. 10, fig. 11.

Ammodochium fletcheri Ling 1985

Plate 1, figures 2-9

Ammodochium fletcheri LING, 1985a, p. 85, pl. 11, figs. 17-20 (not pl. 13, fig. 5).

Remarks: In this species, the proclades and opisthoclades arise from the bifurcated end of the actins. Ammodochium complexum and A. speciosum Deflandre are similar to this taxon, but A. fletcheri has no mesoclades (Ling 1985a). In the double skeletons, two individuals have in common a proclade and opisthoclade skeleton set beside the other flipped skeleton.
Ammodochium sp. cf. A. fletcheri Ling 1985

Plate 1, figure 10
cf. Ammodochium fletcheri LING 1985a, p. 85, pl. 11, figs. 17-20 (not pl. 13, fig. 5).

Remarks: The pore member and alignment on the pro- and opisthoclades differ from those of the typical species. It is possible that this specimen belongs to a new species.
Ammodochium lomonosovense, Onodera and Takahashi sp. nov.

Plate 1, figures 11, 12; Plate 2, figures 1-5, 6?; Plate 4, figure 10
Ammodochium fletcheri LING 1985a, pl. 13, fig. 5 (only).

Jonaotaro Onodera and Kozo Takahashi: Middle Eocene ebridians from the central Arctic Basin

Ammodochium rectangulare (Schulz) Deflandre 1933

Plate 2, figures 7-12

Ebria antiqua rectangularis SCHULZ 1928, p. 274, figs. 72a-72d. Ammodochium prismaticum HOVASSE 1932a, p. 121, figs. 2a-2d. Ammodochium rectangulare (Schulz) DEFLANDRE 1933, p. 517-518, figs. 5-7.

two lateral pores. The rare occurrence of this taxon was recorded in Lithologic Unit 2 in the Core 302-M4A.
Ammodochium sp. 3

Plate 2, figures 16, 17 Remarks: The pro- and opisthoclade look like a wall rather than bar structure. The forms of the pores are different from one another. This rare taxon was observed in Lithologic Unit 2. Genus Ebriopsis Hovasse, 1932a
Ebriopsis crenulata Hovasse, 1932

Ammodochium speciosum Deflandre 1934

Plate 2, figure 14
Ammodochium speciosum DEFLANDRE 1934, p. 92-94, figs. 37, 38.

Remarks: The skeleton of this species has one large pore in each proclade and opisthoclade, and six small pores, framed by proclade, opisthoclade, and mesoclade in the middle of the skeleton.
Ammodochium sp. 1

Plate 3, figure 1
Ebria antiqua SCHULTZ 1928, p. 272, (in part). Ebriopsis crenulata HOVASSE 1932b, p. 281, fig. 4.

Plate 2, figure 13 Remarks: The proclades of this species bifurcate twice giving rise to three pores: one is on the proclade and two at the contact between proclade and two synclades. The posterior half of this species resembles A. rectangulare and the anterior half structure resembles a half of A. complexum Dumitrica & Perch-Nielsen. This taxon is rare in the lower part of Lithologic Unit 2.
Ammodochium sp. 2

Remarks: This taxon was previously reported from upper Eocene and the Oligocene sediments (Schultz 1928; Hovasse, 1932b; Perch-Nielsen, 1976). However, this taxon is usually observed in this study in Cores 302-M4A-8X, 9X, and 10X of early middle Eocene age. Genus Falsebria Deflandre, 1951
Falsebria spp.

Plate 3, figures 2-4

Falsebria ambigua DEFLANDRE 1951, p. 33, 73, figs. 88, 89, 100, 101. Falsebria sp. in PERCH-NIELSEN 1976, p. 153, pl. 10, figs. 1-10.

Plate 2, figure 15 Remarks: This taxon resembles Ammodochium complexum in the general structure, but differs from it in the absence of the

Remarks: The forms of the simple Falsebria skeleton defined by Deflandre (1951) are constituted by the triaene, actins,

Middle Eocene ebridians. All scale bars = 10m. 1 Ammodochium complexum Dumitrica and PerchNielsen, Sample IODP302-M4A-4X-CC, 0-3cm, 2-9 Ammodochium fletcheri Ling 2-7, 8 (Double skeleton), (2-5: IODP302-M2A-61X-CC, 0-1cm; 6: IODP302-M2A-61X-2, 36-38cm; 7: IODP302M4A-6X-1, 36-38cm; 8: IODP302-M2A-58X-2, 96-98cm; 9: IODP302-M2A-61X-1, 56-58cm), 10 Ammodochium sp. cf. fletcheri Ling, IODP302M2A-58X-4, 56-58cm, 11,12 Ammodochium lomonosovense, sp. nov., (11 (Holotype): IODP302-M2A-61X-2, 76-78cm, 12: IODP302-M2A-59X-2, 136-138cm).

PLATE 1

192

Jonaotaro Onodera and Kozo Takahashi

Plate 1

micropaleontology, vol. 55, nos. 2-3, 2009

193

Jonaotaro Onodera and Kozo Takahashi: Middle Eocene ebridians from the central Arctic Basin

rhabde, and rarely an incomplete proclade or synclade. These forms are considered as early ontogenic stages of the ebridians (Perch-Nielsen, 1976; Tappan 1980). Genus Haplohermesinum Hovasse, 1943
Haplohermesinum cornuta (Dumitrica and Perch-Nielsen) Locker 1996

Remarks: The species is previously known to occur in the Miocene (Loeblich et al. 1968). However, this species in this study is sometimes observed throughout Lithologic Units 2 and 1/6 (Appendix 1). It may suggest the extension of occurrence age or the rework. Genus Hermesinopsis Deflandre, 1934
Hermesinopsis valida (Deflandre) Locker 1996

Plate 3, figures 5-11

Ebriopsis cornuta Dumitrica and Perch-Nielsen, in PERCH-NIELSEN 1975, p. 152, pl. 8, figs. 15, 16; pl. 10, fig. 11; LING 1985, p. 85, pl. 11, fig. 24, pl. 13, fig. 4. Haplohermesinum cornuta (Dumitrica and Perch-Nielsen) LOCKER 1996, p. 114, pl. 5, fig. 19.

Plate 4, figures 4-6

Parebria valida DEFLANDRE 1934, p. 91, figs. 33-36. Haplohermesinum validum (Deflandre) HOVASSE 1944, p. 68. Ebriopsis valida (Deflandre) DEFLANDRE 1950, p. 1683. Hermesinopsis valida (Deflandre) LOCKER 1996, p. 114, pl. 5, fig. 25.

Remarks: The hypersilicified skeleton has the external silica rods or silica wall formed around the original skeletal structure in the lorica development process. The abundance of the skeletons with lorica was 5% of all skeletons of Ebriopsis conuta . Genus Hermesinella Deflandre 1934
Hermesinella transversa Deflandre 1934

Genus Hermesinum Zacharias 1906

Hermesinum sp.

Plate 4, figure 7 Remarks: This taxon with lorica occurs in the coarse fraction of Samples 302-M4A-10X-1 and 10X-2. Genus Parebriopsis Hovasse, 1932c
Parebriopsis symmetrica Dumitrica and Perch-Nielsen 1976

Plate 3, figures 12-16

Hermesinella transversa DEFLANDRE 1934, p. 82, 83, figs.14-17.

Hermesinella schulzii (Hovasse) Locker and Martini 1989

Plate 4, figures 1-3

Hermesinum schulzii HOVASSE 1932b, p.125. Hermesinella schulzii (Hovasse) LOCKER and MARTINI 1989, p. 572, pl. 7, fig. 8.

Plate 4, figure 8
Parebriopsis symmetrica Dumitrica and Perch-Nielsen in PERCHNIELSEN 1976, p. 153, pl. 10, figs. 10, 14.

Middle Eocene ebridians. All scale bars = 10m. 1-5,6? (6: Double skeleton?). Ammodochium lomonosovense, sp. nov., (1: IODP302-M4A-4X-1, 0-3cm; 2, 6: IODP302-M2A-61X-CC; 3, 5: IODP302-M2A59X-2, 136-138cm; 4: IODP302-M4A-6X-1, 36-38cm), 7-12 (11, 12: double skeleton). Ammodochium rectangulare (Schulz) Deflandre, (7: IODP302M2A-48X-CC, 4-6cm; 8: IODP302-M2A-53X-1, 36-38cm; 9, 10: IODP302-M2A-48X-4, 14-16cm; 1 1 : IOD P 3 0 2 -M 2 A -4 8 X -C C , 4 -6c m; 12 : IODP302-M2A-53X-2, 14-16cm), 13 Ammodochium sp. 1, IODP302-M2A-61X-2, 76-78cm, 14 Ammodochium speciosum Deflandre, IODP302M2A-49X-4, 14-16cm, 15 Ammodochium sp. 2, IODP302-M4A-11X-3, 128-130cm, 16,17 Ammodochium sp. 3, IODP302-M2A-48X-2, 14-16cm.

PLATE 2

194

Jonaotaro Onodera and Kozo Takahashi

Plate 2

micropaleontology, vol. 55, nos. 2-3, 2009

195

Jonaotaro Onodera and Kozo Takahashi: Middle Eocene ebridians from the central Arctic Basin

Remarks: The occurrence of this taxon is very rare. This taxon was common in the Eocene Norwegian Sea sediments (Perch-Nielsen 1976). Genus Polyebriopsis Hovasse, 1932c
Polyebriopsis sp.

Pseudammodochium karyon, Onodera and Takahashi sp. nov.

Plate 6, figures 1-3 Derivation of Name: The specific name karyon was derived from the Greek word (neuter) meaning walnut. Holotype: Plate 6, figure 3, sample IODP-302-M2A-49X-2, 136-138cm, slide MPC-02688 (Micropaleontology Collection, National Science Museum, Tokyo), England finder J28/4. Type locality: IODP-302 Site M0002 Hole A, 211.51-211.53 mbsf, Lomonosov Ridge in the central Arctic Ocean Description: The outline is ovoid in lateral view. The ring diameter of one end is larger than that of the other end. The outline in apical view is nearly triangular with rounded corners. Actines bifurcated in horizontal plane, their ends reaching to the rounded corners. Clades, arising from the distal ends of actines, represent the scaffolding on which the skeletal wall is built. Pores regularly arranged. Dimensions: length 27-35m; width 24-33m [N = 30]. Occurrence: Cores M2A-50X to 48X with the abundances of <30% of all ebridians. Remarks: This taxon is distinguished from Pseudammodochium dictyoides by its more triangular outline in axial view, slightly larger size than the latter, and regularly arranged pores.
Pseudammodochium sp. cf. karyon, Onodera and Takahashi sp. nov.

Plate 4, figure 9 Genus Pseudammodochium Hovasse, 1932c

Pseudammodochium dictyoides Hovasse 1932

Plate 5, figures 1-10

Pseudammodochium dictyoides HOVASSE 1932c, p. 463, figs. 12-15. Pseudammodochium sp. in PERCH-NIELSEN 1976, p. 154, pl. 6, figs. 17-20.

Remarks: In the ACEX samples, the pores framed by the network of the skeleton are usually fine compared to those of the holotype. The skeletons with coarser pores were rare compared to those having fine pores. The specimens with small pores resemble those of Pseudammodochium sp. in Perch-Nielsen (1976).
Pseudammodochium eximium Deflandre 1951

Plate 5, figures 11, 12

Pseudammodochium eximium DEFLANDRE 1951, p. 56, 80, fig. 213.

Remarks: This species with ovoidal outline differs from Pseudammodochium dictyoides in having almost the same apical and antapical ring diameters.

Plate 6, figures 4, 5

Middle Eocene ebridians. All scale bars = 10m. 1 Ebriopsis crenulata Hovasse, IODP302-M4A-4X-1, 0-3cm, 2-4 Falsebria spp., IODP302-M4A-4X-CC, 0-3cm, 5-11 Haplohermesinum cornuta (Dumitrica and PerchNielsen) Locker, (5: IODP302-M2A-58X-1, 36-38cm; 6: IODP302-M2A-48X-CC, 4-6cm; 7: IODP302-M4A-4X-1, 0-3cm; 8: IODP302-M2A49X-4, 14-16cm; 9, 11: IODP302-M2A-48X-4, 14-16cm; 10: IODP302-M2A-48X-4, 14-16cm), 12-16 Hermesinella transversa Deflandre, (12: IODP302M4A-6X-1, 36-38cm; 13: IODP302-M4A-4X-1, 0-3cm; 14: IODP302-M2A-58X-1, 36-38cm; 15: IODP302-M2A-59X-2, 136-138cm; 16: IODP302M2A-61X-2, 76-78cm).

PLATE 3

196

Jonaotaro Onodera and Kozo Takahashi

Plate 3

micropaleontology, vol. 55, nos. 2-3, 2009

197

Jonaotaro Onodera and Kozo Takahashi: Middle Eocene ebridians from the central Arctic Basin

Remarks: It differs from Pseudammodochium karyon by the absence of two pores on the crest between proclade and opisthoclade.
Pseudammodochium robustum Deflandre 1934

Occurrence: Constant occurrence in Lithologic Unit 2 with abundances below 10%. Remarks: This species differs from other Pseudammodochium species in that the proclades and opisthoclades are relatively broad, and the pores are scarce and small. It is smaller than other ebridian species in the assemblage.
Pseudammodochium sp. 1

Plate 5, figures 13-14

Pseudammodochium robustum DEFLANDRE 1934, p. 94, 95, figs. 39-42.

Remarks: In the ACEX samples, this taxon usually occurred as double skeletons.
Pseudammodochium psichion Onodera and Takahashi sp. nov.

Plate 6, figure 9 Remarks: This species resembles Pseudammodochium karyon, but is distinguished from the latter by smaller and denser pores. This taxon is very rare in Unit 2 and the lower part of Unit 1/6.
Pseudammodochium sp. 2

Plate 6, figures 6-8 Derivation of Name: From psichion, is Greek word (neuter) meaning grain or small drop. Holotype: Plate 6, figure 8, sample IODP-302-M4A-11X-1, 8-10cm, slide MPC-02689 (Micropaleontology Collection, National Science Museum, Tokyo), England finder O22/1. Type locality: IODP Exp. 302 Site M0004 Hole A, 297.38-297.40 mbsf, Lomonosov Ridge in the central Arctic Ocean Description: Skeleton small, ovoid or nearly circular in lateral view, triangular with rounded corners in axial view. The sides of the triangle are concave. Skeleton consisting of essentially a siliceous wall, probably due to heavy silicification. The wall is thick with small pores. Dimensions: length 19-28m; width 18-26m [N = 20].

Plate 4, figure 10 Remarks: In the lateral view, relatively large elliptic pores are observed analogous to those in Ammodochium lomonosovense. However, the number of pores is larger than that of A. lomonosovense. The denser arrangement of pores are characteristic for the genus Pseudammodochium in contrast with that of Ammodochium. The pore arrangement is different from that of P. dictyoides and P. karyon. This taxon was rarely observed in Lithologic Unit 2. Genus Spongebria Deflandre, 1950
Spongebria marthae Deflandre 1950

Plate 6, figures 9-11

Spongebria marthae DEFLANDRE 1950, p. 159, figs. 10, 11.

Middle Eocene ebridians. All scale bars = 10m. 1-3 Hermesinella schulzii (Hovasse) Locker and Martini, (1: IODP302-M2A-61X-2, 76-78cm; 2: IODP302M2A-61X-CC, 0-1cm; 3: IODP302-M4A-6X-1, 14-16cm), 4-6 Hermesinopsis valida (Deflandre) Locker, (4: IODP302-M2A-53X-4, 36-38cm; 5: IODP302M4A-4X-1, 0-3cm; 6: IODP302-M2A-61X-1, 56-58cm), 7 Hermesinum sp. with lorica, IODP302-M4A-10X-2, 56-58cm, 8 Parebriopsis symmetrica Dumitrica and Perch-Nielsen, IODP302-M2A-48X-2, 14-16cm, 9 Polyebriopsis sp., IODP302-M2A-54X-CC, 5-7cm. 10 Double skeleton of Ammodochium lomonosovense, IODP302-M2A-59X-2, 136-138cm.

PLATE 4

198

Jonaotaro Onodera and Kozo Takahashi

Plate 4

micropaleontology, vol. 55, nos. 2-3, 2009

199

Jonaotaro Onodera and Kozo Takahashi: Middle Eocene ebridians from the central Arctic Basin

Remarks: In this species each face of the triode center has a triangular button with rough surface giving an impression of sponge structure by LM observation. Some specimens show that the actines may be connected by synclades, a fact illustrated also by Deflandre (1950). ACKNOWLEDGMENTS We thank the co-chief scientists Professor Jan Backman and Professor Kate Moran, who brought the ACEX cruise to fruition, and the ACEX scientists together with the captains and crew of the IODP Expedition 302 ACEX for their assistance in various phases. We thank Prof. Dimitrios Zafiropoulos of the American University of Athens who assisted us with the new taxonomic names. Drs. Paulian Dumitrica and Katharina von Salis Perch-Nielsen significantly improved this manuscript as pre-reviewers, and their input gratefully appreciated. The SEM photography was performed using a Shimazu SS-550 at the Center of Advanced Instrumental Analysis, Kyushu University. This research was partially supported by a JSPS Research Fellowship to JO as well as JSPS B Project No. 17310009, JSPS B2 No. 10480128, B1 No. 13440152, and B2 No. 15310001 to KT. A part of this research has been supported by Prof. Tatsuro Matsumoto Scholarship Funds of the Kyushu University.
REFERENCES
BACKMAN, J., MORAN, K., MCINROY, D. B., MAYER, L. A. et al., 2006. Proceedings of Integrated Ocean Drilling Program 302. College Station, TX: Integrated Ocean Drilling Program Management International, Inc. doi:10. 2204/iodp. proc. 302. 2006.

BACKMAN, J., JAKOBSSON, M., FRANK, M., SANGIORGI, F., BRINKHUIS, H., STICKLEY, C., OREGAN, M., LVLIE, R., PLIKE, H., SPOFFORTH, D., GATTACECCA, J., MORAN, K., KING, J. and HEIL, C. 2008. Age model and core-seismic integration for the Cenozoic Arctic Coring Expedition sediments from the Lomonosov Ridge. Paleoceanography 23: PA1S03, doi:10. 1029/2007PA001476, 2008. BOHATY, S. M. and HARWOOD, D. M., 2000. Ebridian and silicoflagellate biostratigraphy from Eocene McMurdo Erratics and the Southern Ocean. In: Stilwell, J. D., and Feldmann, R. M., Eds., Paleobiology and paleoenvironments of Eocene rocks, McMurdo Sound, East Antarctica, 99-159. Antarctic Research Series 76. BRINKHUIS, H., SCHOUTEN, S., COLLINSON, M. E., et al., 2006. Episodic fresh surface waters in the Eocene Arctic Ocean. Nature, 441: 606-609. BUKRY, D., 1984. Paleogene paleoceanography of the Arctic Ocean is constrained by the middle or late Eocene age of the USGS core Fl-422; evidence from silicoflagellates. Geology, 12: 199-201. CLARK, D. L., 1974. Late Mesozoic and early Cenozoic sediment cores from the Arctic Ocean. Geology, 2: 41-44. CLARK, D. L., BYERS, C. W. AND PRATT, L. M., 1986. Cretaceous black mud from the central Arctic Ocean. Paleoceanography, 1: 265-271. DEFLANDRE, G., 1933. Enkystement et stade loriqu chez les briaces. Bulletin de la Socit zoologique de France, 57: 514-523. , 1934. Nomenclature du squelette des Ebriaces et description de quelques formes nouvelles. Annales de Protistologie, 4: 75-96.

Middle Eocene ebridians. All scale bars = 10m. 1-10 (6-10: double skeleton). Pseudammodochium dictyoides Hovasse, (1, 6, 9, 10: IODP302M2A-49X-2, 14-16cm; 2: IODP302-61X-CC, 0-1cm; 3: IODP302-M2A-53X-1, 36-38cm; 4: IODP302M2A-48X-CC, 4-6cm; 5: IODP302-M4A-6X-1, 36-38cm; 7: IODP302-M2A-53X-1, 36-38cm; 8: IODP302-M2A-49X-4, 14-16cm), 11,12 Pseudammodochium eximium Deflandre, (11: IODP302-M2A-48X-2, 14-16cm; 12: IODP302M2A-48X-4, 14-16cm), 13,14 Pseudammodochium robustum D e f l a n d r e , IODP302-M2A-49X-4, 14-16cm.

PLATE 5

200

Jonaotaro Onodera and Kozo Takahashi

Plate 5

micropaleontology, vol. 55, nos. 2-3, 2009

201

Jonaotaro Onodera and Kozo Takahashi: Middle Eocene ebridians from the central Arctic Basin

, 1950. Sur une tendance volutive des bridiens. Comptes rendus hebdomadaires des sances de lacadmie des sciences, 231: 158-160. , 1951. Recherches sur les bridiens. Palobiologie. volution. Systmatique. Bulletin Biologique de la france et de la Belgique, 85: 1-84. DELLAGNESE, D. J. and CLARK, D. L., 1994. Siliceous microfossils from the warm Late Cretaceous and Early Cenozoic Arctic Ocean. Journal of Paleontology, 68: 31-47. ERNISSEE, J. J. and MCCARTNEY, K., 1993. Ebridians. In: Lipps, J. H., Ed., Fossil Prokaryotes and Protists, 131-140. London: Blackwell Scientific Publications. GLESER, Z. I., 1996. Problems of Eocene siliceous phytoplankton zonation (exemplified by the Caspian Eocene deposits). Stratigraphy and Geological Correlation, 4: 392-402. HARGRAVES, P. E., 2002. The ebridian flagellates Ebria and Hermesinum. Plankton Biology and Ecology, 49: 9-16. HOVASSE, R., 1932a. Note Prliminaire sur les briaces. Bulletin de la Socit zoologique de France, 57: 118-131. , 1932b. Seconde note sur les briaces. Bulletin de la Socit zoologique de France, 57: 278-283. , 1932c. Troisime note sur les briaces. Bulletin de la Socit zoologique de France, 57: 457-476. , 1943. Nouvelles recherches sur les flagells squelette siliceux: briids et silicoflagells fossiles de la diatomite de

Saint-Laurent-La-Vernde (Gard). Bulletin Biologique de la france et de la Belgique,77: 285-294. , 1944. propos des silicoflagells fossils. Rplique M. G. Deflandre. Bulletin Biologique de la france et de la Belgique, 78: 68-69. KITCHELL, J. A. and CLARK, D. L., 1982. Late Cretaceous-Paleogene paleogeography and palercirculation: Evidence of north polar upwelling. Palaeogeography, Palaeoclimatology, Palaeoecology, 40: 135-163. KORHOLA, A. and SMOL, J. P. 2001. Ebridians. In: Smol, J. P., Birks, H. J. B. and W. M. Last, Eds., Tracking Environmental Change Using Lake Sediments Volume 3: Terrestrial, Algal, and Siliceous Indicators, 225-234. Dordrecht: Kluwer Academic Press. LING, H. Y., 1972. Upper Cretaceous and Cenozoic silicoflagellates and ebridians. Bulletins of American Paleontology, 62: 135-229. , 1985a. Early Paleogene silicoflagellates and ebridians from the Arctic Ocean. Transaction Proceedings of the Palaeontological Society of Japan, New Series, 138: 79-93. , 1985b. Paleogene silicoflagellates and ebridians from the Goban Spur, northeastern Atlantic. In: Graciansky, P. C. de, Poag, C. W., et al., Initial Reports of Deep Sea Drilling Project 80, 663-668. Washington, DC: U. S. Government Printing Office. LOCKER, S., 1996. Cenozoic siliceous flagellates from the Fram Strait and the East Greenland Margin: biostratigraphic and paleoceanographic results. In: Thiede, J, Myhre, A. M., Firth, J. V., Johnson, G. L. and Ruddiman, W. F., Eds., Proceedings of the Ocean Drilling Program, Scientific Results 151, 101-124. College Station, TX: Ocean Drilling Program.

Middle Eocene ebridians. All scale bars = 10m. 1-3 Pseudammodochium karyon, sp . n o v . , ( 1 : IODP302-M2A-48X-2, 14-16cm; 2: IODP302M2A-48X-4, 14-16cm; 3 (Holotype): IODP302M2A-49X-2, 136-138cm), 4,5 Pseudammodochium sp. cf. karyon, sp. nov., (4: IODP302-M2A-48X-2, 14-16cm; 5: IODP302M4A-11X-3, 128-130cm) 6-8 Pseudammodochium psichion sp. nov., (6: IODP302-M2A-48X-CC, 4-6cm; 7: IODP302M2A-59X-2, 136-138cm; 8 (Holotype): IODP302M4A-11X-1, 8-10cm), 9 Pseudammodochium sp. 1, IODP302-M2A-48X-CC, 4-6cm, 10-12 Spongebria marthae Deflandre, (5: IODP302M2A-58X-1, 36-38cm; 6: IODP302-M4A-4X-1, 0-3cm; 7: IODP302-M2A-48X-CC, 4-6cm).

PLATE 6

202

Jonaotaro Onodera and Kozo Takahashi

Plate 6

micropaleontology, vol. 55, nos. 2-3, 2009

203

Jonaotaro Onodera and Kozo Takahashi: Middle Eocene ebridians from the central Arctic Basin

LOCKER, S. and MARTINI, E., 1986. Silicoflagellates and some sponge spicules from the southwest Pacific, Deep Sea Drilling Project, Leg 90. In: Kennet, J. P., von der Borch, C. C., et al., Initial Reports of the Deep Sea Drilling Project 90, 887-924. Washington, DC: U. S. Government Printing Office. LOEBLICH, A. R. III., LOEBLICH, L. A., TAPPAN, H. AND LOEBLICH, A. R. Jr., 1968. Annotated index of fossil and recent silicoflagellates and ebridians with descriptions and illustrations of validly proposed taxa. Boulder, CO: Geological Society of America. Memoir 106, 319 pp. MAGAVERN, S., CLARK, D. L. AND CLARK, S. L., 1996. 87/86Sr, phytoplankton, and the nature of the Late Cretaceous and Early Cenozoic Arctic Ocean. Marine Geology 133: 183-192. MORAN, K., BACKMAN, J., BRINKHUIS, H. et al., 2006. The Cenozoic palaeoenvironment of the Arctic Ocean. Nature, 441: 601-605. ONODERA, J. and TAKAHASHI, K, 2009. Taxonomy and biostratigraphy of silicoflagellates in the middle Eocene Arctic Ocean. Micropaleontology, this volume. ONODERA, J., TAKAHASHI, K. and JORDAN, R. W., 2008. The Eocene silicoflagellate and ebridian paleoceanography in the central Arctic Ocean. Paleoceanography, 23, PA1S15, doi:10. 1029/2007PA001474. OSAWA, M., TAKAHASHI, K. and HAY, B. J., 2005. Shell-bearing plankton fluxes in the central Black sea, 1989-1991. Deep-Sea Research I, 52: 1677-1698.

PERCH-NIELSEN, K., 1976. Eocene to Pliocene archaeomonads, ebridians, and endoskeletal dinoflagellates from the Norwegian Sea, DSDP Leg 38. In: Talwani, M., Udintsev, G., et al., Initial Reports of the Deep Sea Drilling Project 38, suppl., 147-175. Washington, DC: U. S. Government Printing Office. SANGIORGI, F., BRINKHUIS, H. and DAMASSA, S. P., 2009. Frigusphaera gen. nov.: A new organic-walled dinoflagellate cyst genus from the ?early Miocene of the central Arctic Ocean. Micropaleontology, this volume. SCHULZ, P., 1928. Beitrge zur Kenntnis fossiler und rezenter Silicoflagellaten. Botanisches Archiv, 21: 225-292. SIMS, P. A. and ROSS, R., 1988. Some Cretaceous and Palaeogene Trinacria (diatom) species. Bulletin of the British Museum of Natural History, 18: 275-322. STICKLEY, C. E., KO, N., BRUMSACK, H., JORDAN, R. W. and SUTO, I. 2008. A siliceous microfossil view of Middle Eocene Arctic palaeoenvironments: a window of biosilica production and preservation. Paleoceanography, 23, PA1S14, doi:10. 1029/2007PA00 1485. TAPPAN, H., 1980. Ebridians. In: Tappan, H., Ed., The paleobiology of plant protists, 463-489. New York: W. H. Freeman and Company. ZACHARIAS, O., 1906. Ene neue Dicyochide aus dem Mittelmeer, Hermesinum adriaticum n. g., n. sp. Archiv fr Hydrobiologie und Planktonkunde, 1: 394-398.

Middle Eocene ebridians. All scale bars = 10m. 1-8 Ebridian gen. et. sp. indet. sp. 1 (1: IODP302M4A-4X-1, 0-3cm; 2: IODP302-M2A-48X-4, 14-16cm; 3-5, 7, 8: IODP302-M2A-48X-4, 14-16cm; 6: Sample 302-M2A-48X-CC, 4-6cm), 9 Ebridian gen. et. sp. indet. sp. 2, Sample 302-M2A48X-4, 14-16cm, 10-12 Ebridian gen. et. sp. indet. sp. 3, (10: IODP302M2A-48X-CC, 4-6cm; 11: IODP30-4X-1, 4-6cm).

PLATE 7

204

Jonaotaro Onodera and Kozo Takahashi

Plate 7

micropaleontology, vol. 55, nos. 2-3, 2009

205