The development of planktonic biogeography in the Southern Ocean during the Cenozoic

Deep-sea drilling at high latitudes of the Southern Hemispheres has provided almost the only available data to evaluate the biogeographic development of the planktonic biota in the Southern Ocean during the Cenozoic (65 m.y. to Present Day). Paleontological investigations on Deep Sea Drilling Project (DSDP) materials have shown that the development of Cenozoic planktonic biogeography of the Southern Ocean is intimately linked with the evolution of the Southern Ocean water masses themselves. During the Cenozoic, this has included the development of the Circum-Antarctic Current system as obstructing land masses moved apart, the refrigeration and later extensive glaciation of the continent, and the development of the Antarctic Convergence (Polar Front) with related oceanic upwelling.Almost all evolution of calcareous planktonic microfossils has occurred outside of the Antarctic—Subantarctic region followed by limited migration into these water masses. Virtually no endemism occurs amongst calcareous microfossil groups at these latitudes. In contrast, conspicuous and widespread evolution has occurred within the siliceous microfossil groups especially during the Neogene. Low diversity and differences in stratigraphic ranges of Antarctic calcareous microfossils makes them only broadly useful for correlation. Relatively higher diversities within the Subantarctic provide a firmer basis for more detailed correlation, although the ranges of fossils are often different than at lower latitudes because of different paleoceanographic and paleoclimatic controls. Within the Antarctic water mass south of the Antarctic Convergence, siliceous microfossilsbiostratigraphy, oxygen isotopic stratigraphy and magnetostratigraphy, provide the only firm basis for correlation with low-latitude sequences.Eocene (55-38 Ma) sediments contain abundant calcareous microfossils even closely adjacent to the continent. Antarctic calcareous planktonic microfossils of this age exhibit relative high diversity, although this is lower than assemblages of equivalent age at middle and low latitudes. Within the Subantarctic region, Eocene planktonic foraminifera exhibit strong affinities with those in the temperate regions. Biogeographic differences exist between various sectors of the Southern Ocean related to biogeographic isolation preceding the development of the Circum-Antarctic Current. Subantarctic calcareous nannofossil assemblages of Paleocene and Eocene age exhibit higher diversity than Oligocene and Neogene assemblages. Siliceous microfossils are poorly represented or at best poorly known.One of the most dramatic changes in Southern Ocean planktonic biogeography occurred near the Eocene/Oligocene boundary (38 Ma). Since then, Antarctic planktonic foraminiferal assemblages have exhibited distinct polar characteristics, marked in particular by low diversity, and this event thus reflects the initiation of the Antarctic faunal and floral provinces. Profound paleoceanographic changes at this time, which triggered the biogeographic crisis, appear to be related to the initiation of widespread Antarctic sea-ice formation, and rapid cooling of deep and intermediate waters, in turn associated with increased Antarctic glaciation. During the Oligocene, planktonic microfossil diversity was low in all groups throughout the world's oceans. In Antarctic waters, the early Oligocene foraminiferal fauna is monospecific (Subbotina angiporoides), while in the later Oligocene two species (S. angiporoides and Catapsydrax dissimilis) were recorded. Calcareous nannofossil assemblages are of low diversity compared with the Eocene. Subantarctic foraminiferal faunas of Oligocene age display much higher diversity than those in the Antarctic, but early and middle Oligoceae faunas still exhibit the lowest diversities for the entire Cenozoic. Siliceous assemblages remain relatively inconspicuous in most regions of the Southern Ocean.The Paleogene-Neogene transition (22 Ma) is marked by a major change in the global planktonic biogeography, i.e. modern patterns developed in which permanent, steep faunal and floral diversity gradients existed between tropical and polar regions; a gradient which has persisted even during the most severe glacial episodes. Oligocene assemblages of low diversity and almost cosmopolitan distribution were replaced by distinctive belts of planktonic assemblages arranged latitudinally from the tropics to the poles. The establishment of the steep planktonic diversity gradients and latitudinal provinces near the beginning of the Neogene almost certainly were linked to the development of the Circum-Antarctic Current in the late Oligocene which effectively separated high- and low-latitude planktonic assemblages. These fundamental global circulation and biogeographic patterns have persisted through the Neogene.During the Neogene (22 Ma to Present Day), Antarctic calcareous microfossil assemblages exhibit persistent low diversity and high dominance, while Subantarctic assemblages are of much greater diversity. The beginning of the Neogene (= beginning of Miocene) heralded the development of the high-latitude siliceous microfossil assemblages towards their present-day dominant role. Siliceous biogenec productivity began to increase. These changes were linked to the initial development and later intensification of circulation associated with the Antarctic Convergence and Antarctic Divergence. The Antarctic Convergence sharply separates dominantly siliceous assemblages to the south from calcareous assemblages to the north. Radiolarian assemblages became more endemic. Relatively warm early and middle Miocene conditions are reflected by slightly higher diversity of planktonic foraminifera and by the presence, in the northern Subantarctic, of conspicuous discoasters in early Miocene sediments. In Antarctic waters, calcareous nannofossils become unimportant as biogenic elements after the middle Miocene.The latest Miocene ( 5 m.y. ago) was marked by northward movement of the Antarctic Convergence, corresponding expansion of the Antarctic water mass, and low diversity of calcareous assemblages. Pliocene planktonic foraminifera seem to be largely monospecific in Antarctic and southern Subantarctic sequences. During the Quaternary, Antarctic waters reached a maximum northward expansion and exhibit highest siliceous biogenic productivity for the Cenozoic. In the Subantarctic, Quaternary foraminiferal diversities are much higher than in Pliocene sequences. Although calcareous nannofossil diversity may be high, only a few species are abundant. Large northward shifts of Antarctic and Subantarctic water masses have occurred during the Quaternary although no southward penetrations have occurred much beyond that of the present day. Several radiolarian and foraminiferal species disappeared or appeared at or close to a number of paleomagnetic reversals during the last 4 m.y. These faunal events, which provide valuable datums, do not seem to be associated with major climatic changes.     

Response of deep-sea benthic foraminifera to Miocene paleoclimatic events, DSDP Site 289

Changes in the Miocene deep-sea benthic foraminifera at DSDP Site 289 closely correlate to the climatically induced variations in deep and bottom waters in the Pacific Ocean. In early Miocene time, oxygen and carbon isotopes indicate that bottom waters were relatively warm and poorly oxygenated. Benthic foraminiferal assemblages are characterized by various species inherited from the Oligocene. Expansion of the Antarctic icecap in the early middle Miocene, 14–16 m.y. ago, increased oxygen isotope values, produced cold, more oxygenated bottom waters and lead to a turnover in the benthic foraminifera. An Oligocene—early Miocene assemblage was replaced by a cibicidoid-dominated assemblage. Some species became extinct and benthic faunas became more bathymetrically restricted with the increased stratification of deep waters in the ocean. In mid-Miocene time, Epistominella exigua and E. umbonifera, indicative of young, oxygenated bottom waters, are relatively common at DSDP Site 289. Further glacial expansion 5–9 m.y. ago lowered sealevel, increased oceanic upwelling and associated biological productivity and intensified the oxygen minima. Abundant hispid and costate uvigerines become a dominant faunal element at shallow depths above 2500 m as E. umbonifera becomes common to abundant below 2500 m. By late Miocene time, benthic faunas similar in species composition and proportion to modern faunas on the Ontong-Java plateau, had become established.     

Cenozoic radiolarian paleobiogeography: Implications concerning plate tectonics and climatic cycles

A preliminary study of the paleobiogeographic patterns of radiolarian facies during the Paleogene and subsequent time shows that:(1) Through time radiolarian assemblages display distinct faunal provincialism reminiscent of modern faunal distributions correlated with planetary temperature gradients and surface oceanic conditions. The equatorial—tropical radiolarian fauna extended apparently unrestricted across the Pacific Ocean, the Caribbean Sea and the Atlantic Ocean through Early Miocene time. In the Caribbean Sea and the Atlantic Ocean, radiolarians reached their maximum abundance in the Eocene and Oligocene. Subsequently, they gradually declined to virtual disappearance in these areas in the early Miocene. Their Pacific counterparts remained practically undisturbed, except that post early Miocene assemblages there showed a marked trend toward decreasing test thickness. This trend has since been a worldwide characteristic of Neogene radiolarian assemblages and their modern equivalents. It is postulated that the disappearance of radiolarians in the Carribean Sea and the Atlantic Ocean at the end of the Paleogene is related to the onset of the emergence of the isthmus of Panama which interrupted the preexisting oceanic circulation between the Pacific and Atlantic Oceans.(2) Throughout the Paleogene there have been marked sequential fluctuations in the radiolarian assemblages of the Caribbean Sea which indicate intermittent incursions of higher-latitude fauna in this area. Associated with the faunal fluctuations are cyclic variations in the total carbonate of the sediment with patterns also comparable in duration to Pleistocene carbonate cycles in the equatorial Pacific known to have been induced by climatic changes. Based on similarities with Pleistocene climatic cycles in the equatorial Pacific and elsewhere, it is surmised that the faunal and lithologic fluctuations observed in Paleogene radiolarian sediments were also induced by the biologic and physico-chemical processes associated with worldwide changes in the climatic conditions of that time.     

Paleocene-Pleistocene benthic foraminiferal evidence of major paleoceanographic events in the eastern South Atlantic (DSDP Site 525, Walvis Ridge)

Benthic foraminifers in the size-fraction greater than 0.073 mm were studied in 88 Paleocene to Pleistocene samples from Deep Sea Drilling Project Site 525 (Hole 525A, Walvis Ridge, eastern south Atlantic). Clustering of the samples on the basis of the 86 most abundant foraminifers (in total, 331 taxa were identified) allowed separating two major assemblage zones: the Paleocene to Eocene interval, and the Oligocene to Pleistocene interval. Each of these, in turn, were subdivided into three minor subzones as follows: lower upper Paleocene (approx. 62.4 to 57.8 Ma); upper upper Paleocene (56.6 to 56.2 Ma); lower and middle Eocene (55.3 to 46.8 Ma); upper Oligocene to middle Miocene (25.3 to 16 Ma); middle Miocene to Pliocene (15.7 to 4.2 Ma); and lower Pleistocene (0.4 to 0.02 Ma), with only minor differences with the previous zone. Some very abundant taxa span most of the column studies (Bolivina huneri, Cassidulina subglobosa, Eponides bradyi, E. weddellensis, Gavelinella micra, Oridorsalis umbonatus, etc.). Several of the faunal breaks recorded coincide with conspicuous minima in the specific diversity curve, thus suggesting that the corresponding turnovers signal the final stages of periods of faunal impoverishment. At least one major bottom-water temperature drop (as derived from δ¹⁸O data) is synchronous with a decrease in the foraminiferal specific diversity. On the other hand, a specific diversity maximum in the middle Miocene might be associated with a δ¹³C increase at approx. 16 to 12 Ma. Highest foraminiferal abundances (up to 600–800 individuals per gram of dry sediment) occurred in the late Paleocene and in the early Pleistocene, in coincidence with the lowest diversity figures calculated. The magnitude of the most important faunal turnover recorded, between the middle Eocene and the late Oligocene, is magnified in our data set by the large hiatus which separates the middle Eocene from the upper Oligocene sediments. Considerably smaller overturns occurred within the late Paleocene (in coincidence with changes in the specific diversity, absolute abundance of foraminiferal tests, and δ¹³C), and in the middle Miocene (in coincidence with a specific diversity maximum and a δ¹³C excursion). New information on the morphology and the stratigraphic ranges of several species is furnished. For all the taxa recorded the number of occurrences, total number of individuals identified and first and last appearances are listed.     

Miocene biochronology and paleoceanography of the North Pacific

Biostratigraphic correlation based on microfossil datum levels, directly or indirectly tied to the paleomagnetic time scale, provides a high resolution time control for the Miocene in the equatorial and middle latitude North Pacific. Faunal changes and abundance fluctuations of planktic foraminiferal species combined with the oxygen Pacific. Faunal changes and abundance fluctuations of planktic foraminiferal species combined with the oxygen isotope record of foraminifers, reveal the paleoclimatic and paleoceanographic history. The planktic foraminiferal assemblage change in the early Miocene, extinction of Oligocene fauna and rise of a highly diverse Neogene fauna, appears to be related to increased water mass stratification in the world oceans presumably resulting from the establishment of circum-Antarctic circulation. An increase in the siliceous productivity in the eastern equatorial Pacific region between 20 and 18 Ma suggests that the vertical and horizontal circulation was intensified at that time. Climates cooled rapidly during the middle Miocene between 14 and 13 Ma suggesting the growth of a major east Antarctic ice sheet. Paleoclimatic conditions remained generally cool, although oscillating, during the late Miocene. In the late early to middle Miocene faunal provincialism developed between low and middle latitudes, and by late Miocene time a distinct provincialism similar to the present was established.     

Latest Cretaceous to Late Paleocene radiolarian biostratigraphy: A new zonation from the New Zealand region

The scarcity of records of Early Paleocene radiolarians has meant that while radiolarian biostratigraphy is firmly established as an important tool for correlation, there has been a long-standing gap between established zonations for the Cretaceous and from latest Paleocene to Recent. It has also led to considerable speculation over the level of faunal change across the Cretaceous/Tertiary (K/T) boundary. Consequently, the discovery of rich and diverse radiolarian assemblages in well-delineated K/T boundary sections within siliceous limestones of the Amuri Limestone Group in eastern Marlborough, New Zealand, is of great significance for biostratigraphy and K/T boundary research.This initial report is restricted to introducing a new latest Cretaceous to mid Late Paleocene zonation based on the radiolarian succession at four of these sections and a re-examination of faunas from coeval sediments at DSDP Site 208 (Lord Howe Rise). Three new Paleocene species are described:Amphisphaera aotea, Amphisphaera kina andStichomitra wero. Six new interval zones are defined by the first appearances of the nominate species. In ascending order these are:Lithomelissa? hoplites Foreman (Zone RK9, Cretaceous),Amphisphaera aotea n. sp. (Zone RP1, Paleocene),Amphisphaera kina n. sp. (RP2),Stichomitra granulata Petrushevskaya (RP3),Buryella foremanae Petrushevskaya (RP4) andBuryella tetradica (RP5). Good age control from foraminifera and calcareous nannofossils permits close correlation with established microfossil zonations. Where age control is less reliable, radiolarian events are used to substantially improve correlation between the sections.No evidence is found for mass extinction of radiolarians at the end of the Cretaceous. However, the K/T boundary does mark a change from nassellarian to spumellarian dominance, due to a sudden influx of actinommids, which effectively reduces the relative abundance of many Cretaceous survivors. An accompanying influx of diatoms in the basal Paleocene of Marlborough, together with evidence for an increase of total radiolarian abundance, suggests siliceous plankton productivity increased across the K/T boundary. Possible causes for this apparently localised phenomenon are briefly discussed.     

Small mammal (rodents and lagomorphs) European biogeography from the Late Oligocene to the mid Pliocene

Aim To analyse the fossil species assemblages of rodents and lagomorphs from the European Neogene in order to assess what factors control small mammal biogeography at a deep‐time evolutionary time‐scale. Location Western Europe: 626 fossil‐bearing localities located within 31 regions and distributed among 18 successive biochronological units ranging from c. 27 Ma (million years ago; Late Oligocene) to c. 3 Ma (mid Pliocene). Methods Taxonomically homogenized pooled regional assemblages are compared using the Raup and Crick index of faunal similarity; then, the inferred similarity matrices are visualized as neighbour‐joining trees and by projecting the statistically significant interregional similarities and dissimilarities onto palaeogeographical maps. The inferred biogeographical patterns are analysed and discussed in the light of known palaeogeographical and palaeoclimatic events. Results Successive time intervals with distinct biogeographical contexts are identified. Prior to c. 18 Ma (Late Oligocene and Early Miocene), a relative faunal homogeneity (high interregional connectivity) is observed all over Europe, a time when major geographical barriers and a weak climatic gradient are known. Then, from the beginning of the Middle Miocene onwards, the biogeography is marked by a significant decrease in interregional faunal affinities which matches a drastic global climatic degradation and leads, in the Late Miocene (c. 11 Ma), to a marked latitudinal pattern of small mammal distribution. In spite of a short rehomogenization around the Miocene/Pliocene boundary (6–4 Ma), the biogeography of small mammals in the mid Pliocene (c. 3 Ma) finally closely reflects the extant situation. Main conclusions The resulting biogeographical evolutionary scheme indicates that the extant endemic situation has deep historical roots corresponding to global tectonic and climatic events acting as primary drivers of long‐term changes. The correlation of biogeographical events with climatic changes emphasizes the prevalent role of the climate over geography in generating heterogeneous biogeographical patterns at the continental scale.     

Foraminiferal paleoecology of the Montpelier and lower coastal groups (eocene-miocene), Jamaica,West Indies

Two domains of carbonate deposition characterized mid-Tertiary Jamaica. After latest Cretaceous to Paleocene orogeny, submergence of insular paleo-Jamaica accompanied the strike-slip or extensional faulting associated with the formation of the Cayman Trench to the north. Differential subsidence along a series of peripheral subsea escarpments (Duanvale-Wagwater escarpment) produced relief in excess of 2000 m by the Late Eocene. Shoalwater limestones covered the slowly subsiding Cornwall-Middlesex platform, thus ending the supply of clastics to deep-sea bottoms north and east of the escarpment where contemporaneous planktonic-foraminiferal pelagites accumulated. Middle Eocene to Middle Miocene carbonate rocks deposited in the deep-sea represent a distinctive lithogenetic unit termed the Montpelier Group.A preponderance of globigerinacean and radiolarian tests characterizes lower Montpelier microfossil assemblages. Dominant benthonic forms include Melonis pompilioides, Fontbotia wuellerstorfi and species of Stilostomella and Pleurostomella. Available faunal criteria including the assemblage composition, depth preferences of extant species and recurrent morphologic-ecologic patterns suggest abyssal (2000 m) paleo-depths at the site of accumulation on a sediment apron near the base of the Duanvale-Wagwater escarpment. Computed from inferred paleodepth and estimated sedimentary thickness, Middle Eocene to Lower Miocene subsidence totals 2800 m. Biostratigraphic and paleoecologic data do not support the prevalent concept of a regional unconformity within the Montpelier.In the Middle Miocene, regional uplift led to the emergence of the Cornwall-Middlesex platform and to pronounced shoaling of marginal sea bottoms. Here, hemipelagic sedimentation resumed during the later Middle Miocene after the carbonate veneer on the adjacent platform was sufficiently eroded so as to expose noncarbonate rocks. See NAPS document No. 02395 for 67 pages of faunal reference lists, maps of sample locations and tables of foraminiferal occurrences. Order from ASIS/NAPS, c/o Microfiche Publications, 305 East 46th St., New York, N.Y., U.S.A. 10017. Remit with order $9.05 for photocopies or $5.55 for microfiche. Make checks payable to Microfiche Publications.     

Biostratigraphy and paleoceanographic significance of Paleocene radiolarians from offshore eastern New Zealand

A 100-m-thick Paleocene sequence of mainly pelagic sediments at ODP Site 1121, on the eastern flanks of the Campbell Plateau, contains few to common radiolarians of relatively low diversity in the lower 40 m (Early to early Late Paleocene) and abundant, diverse radiolarian assemblages in the upper 60 m (mid-Late Paleocene). The 150 taxa recorded from the entire Paleocene interval are thought to under-represent the actual species diversity by at least one half as many morphotypes have not been differentiated below the level of genus. Assemblages in the lower 40 m are similar to those described from onland New Zealand and DSDP Site 208 (northern Lord Howe Rise); they are correlated with South Pacific radiolarian zones RP4 and RP5. Assemblages in the upper 60 m differ from other known Late Paleocene assemblages in the great abundance of plagiacanthids and cycladophorids. Similarities are noted with later Cenozoic cool-water assemblages. This upper interval is correlated with South Pacific zone RP6, as revised herein, based on comparison with faunas from Site 208 and Marlborough, New Zealand. The interval is also correlated with the upper part of North Atlantic zone RP6 (RP6b–c) based on the presence of Aspis velutochlamydosaurus, Plectodiscus circularis and Pterocodon poculum. Other species, such as Buryella tetradica and Buryella pentadica, are valuable for local correlation but exhibit considerable diachroneity between the Pacific, Indian and Atlantic Oceans. An age model for the Paleocene interval at Site 1121, based on well-constrained nannofossil and radiolarian datums, indicates that the rate of compacted sediment accumulation doubles from 15 to 30 mm/ka at the RP5/RP6 zonal boundary. In large part this is due to a sudden and pronounced increase in accumulation rates for all siliceous fossils; radiolarians and larger diatoms increase from <100 to >10 000 specimens/cm²/ka. This apparent increase in biosiliceous productivity is age-equivalent to a mid-Paleocene cooling event (57–59 Ma) identified from global stable isotope records that is associated with the heaviest δ¹³C values for the entire Cenozoic.     

Comparison of radiolarian and sedimentologic paleoproductivity proxies in the latest Miocene–Recent Benguela Upwelling System

Estimating past ocean productivity from ocean sediments often gives different results depending on the measurement used. We have examined a suite of paleoproductivity proxies in latest Miocene–Recent sediments from DSDP Site 532 and ODP Site 1084, two deep-sea sections underlying the Benguela Upwelling System off the Atlantic coast of southern Africa. The productivity history of this system has been previously established via organic carbon concentration, diatom floras and alkenone based estimates of surface water temperature, and shows a change from low productivity in the early Pliocene to sustain high productivity in the late Pliocene–Recent. Each of our samples was split and simultaneously analysed for several proxies of ocean productivity, including organic carbon (TOC%), carbonate, abundance of opaline radiolarians, accumulation rate of benthic foraminifera (BFAR); the radiolarian faunal composition indices Upwelling Radiolarian Index (URI) and the Water Depth Ecology index (WADE); other proxies for opal and carbonate dissolution, plus stable isotopes of benthic foraminifera. Comparisons between proxies in the same measured samples, between sites in downcore plots and to the published productivity record for this region suggest that TOC and radiolarian faunal composition, particularly the WADE index, are good indicators of past productivity, albeit with different sensitivities (log–linear correlation WADE–TOC% r = 0.78, n = 65, p < 0.01). In contrast, carbonate, and carbonate-based proxies such as BFAR primarily reflect changes in dissolution. Radiolarian faunal composition indices do not appear to be affected by bulk opal accumulation or changes in opal preservation. WADE analysis of radiolarian faunas and TOC% measurements appear to be useful proxies for productivity in late Neogene sediments, particularly for sections where opal or carbonate dissolution is significant.     

The Middle Miocene climatic transition in the Southern Ocean: Evidence of paleoclimatic and hydrographic changes at Kerguelen plateau from planktonic foraminifers and stable isotopes

Middle Miocene (14.8–11.9 Ma) deep-sea sediments from ODP Hole 747A (Kerguelen Plateau, southern Indian Ocean) contain abundant, well-preserved and diverse planktonic foraminiferal assemblages. A detailed study of the climatic and hydrographic changes that occurred in this region during the Middle Miocene Climatic Transition led to the identification of an intense cooling phase (the Middle Miocene Shift). Abundance fluctuations of planktonic foraminiferal species with different paleoclimatic affinities, and oxygen and carbon stable isotopes have been integrated in a multi-proxy approach. Reconstruction of changes in foraminiferal faunal composition and diversity through time were the basis for identification of three foraminiferal biofacies. The most prominent faunal change took place at 13.8 Ma, when a fauna with warm-water affinity (marked by high abundance of Globorotalia miozea group and Globoturborotalita woodi plexus) was replaced by an oligotypic, opportunistic fauna with typical polar characters and dominated by neogloboquadrinids. This faunal change is interpreted as the result of foraminiferal migration from adjacent bioprovinces, caused by modifications in climate and hydrography. A positive 2.0‰ shift in δ¹⁸O (interpreted as the Mi3 event) and a related positive 1.0‰ shift in δ¹³C (corresponding to the CM6 event) accompanied this faunal turnover. These are interpreted to reflect substantial reorganization of Southern Ocean waters, the northward migration of the Polar Front and a strong increase in primary productivity. The second faunal change took place at 12.9 Ma and was characterized by the gradual decrease in abundance of the neogloboquadrinids and the recovery of Globorotalia praescitula/scitula group and Globigerinita glutinata. A positive 1.5‰ shift in δ¹⁸O (interpreted as the Mi4 event) and a concurrent gradual negative shift in δ¹³C accompanied this faunal change, witnessing further modifications of the climate/ocean system. Variations in sea surface temperature, considered as the main factor causing changes of surface hydrography at the Kerguelen Plateau, seem to have been driven by obliquity and long-term eccentricity, thus suggesting a key role played by the astronomical forcing on the evolution of Southern Ocean dynamics during the Middle Miocene. Also an evident 1.2 Myr modulation of the δ¹³C record suggests a main control of the long-term obliquity cycles on the carbon cycle dynamics. Particularly, the Mi3/CM6 events exactly fit with a node of the 1.2 Myr modulation cycles. This confirms the key role played by orbital parameters on high-latitude temperatures and Antarctic ice volume, and indirectly on global carbon burial and/or productivity. This climatic transition was marked also by changes in surface hydrography. From 14.8 to 13.8 Ma an intermediate-strength thermocline controlled by seasonality developed just below the photic zone. Weaker seasonality characterized the interval from 13.8 to 12.9 Ma, when the thermocline became shallower and sharper and favored intermediate-water foraminifers. From 12.9 Ma, seasonality increased again and an intermediate-strength thermocline re-developed.     

Density and assemblage influence the nature of the species richness–productivity relationship in Australian dry sclerophyll forest species

The relationship between species richness and productivity is important from both a basic, theoretical perspective and also because it has important ramifications for applied ecology including ecosystem restoration and the design of carbon offset plantings. While a more species‐rich community is often believed to be more productive than a species‐poor community, findings from observational and experimental studies differ and our understanding of the relationship comes largely from grasslands. Consequently, we aimed to determine for the first time the nature of the species richness–productivity relationship in a southern‐hemisphere dry sclerophyll ecosystem. We investigated the impact of species richness on productivity, plant density and mean plant biomass at three sowing densities in three species assemblages. Eucalyptus globulus, Acacia mearnsii and Allocasuarina verticillata were each grown as monocultures and included in every subsequent level of species richness, forming three distinct species assemblages. Communities were grown in a glasshouse pot experiment for four months, then harvested and above‐ground biomass measured. We found no general species richness–productivity relationship in the communities studied. There were no overall increases in productivity as species richness increased and in fact in most cases the productivity of communities with 4 and 8 species was lower than monocultures of the dominants. Importantly, density influenced the way richness affected productivity and this effect was dependent upon assemblage, indicating that species identity is a key determinant of productivity. These results demonstrate important ecological principles in a previously untested system. A key outcome of this experiment is that density alters the relationship between species richness and initial productivity in assemblages of Australian dry sclerophyll species.     

Neogene reef coral assemblages of the Bocas del Toro region, Panama: the rise of Acropora palmata

Temporal patterns are evaluated in Neogene reef coral assemblages from the Bocas del Toro Basin of Panama in order to understand how reef ecosystems respond to long-term environmental change. Analyses are based on a total of 1,702 zooxanthellate coral specimens collected from six coral-bearing units ranging in age from the earliest Late Miocene to the Early Pleistocene: (1) Valiente Formation (12–11 Ma), (2) Fish Hole Member of the Old Bank Formation (5.8–5.6 Ma), (3) La Gruta Member of the Isla Colon Formation (2.2–1.4 Ma), (4) Ground Creek Member of the Isla Colon Formation (2.2–1.4 Ma), (5) Mimitimbi Member of the Urracá Formation (1.2–0.8 Ma), and (6) Hill Point Member of the Urracá Formation (1.2–0.8 Ma). Over 100 coral species occur in the six units, with faunal assemblages ranging from less than 10% extant taxa (Valiente Formation) to over 85% extant taxa (Ground Creek Member). The collections provide new temporal constraints on the emergence of modern Caribbean reefs, with the La Gruta Member containing the earliest occurrence of large monospecific stands of the dominant Caribbean reef coral Acropora palmata, and the Urracá Formation containing the last fossil occurrences of 15 regionally extinct taxa. Canonical correspondence analysis of 41 Late Miocene to Recent reef coral assemblages from the Caribbean region suggests changes in community structure coincident with effective oceanic closure of the Central American Seaway (~3.5 Ma). These changes, including increased Acropora dominance, may have contributed to a protracted period of elevated extinction debt prior to the major peak in regional coral extinctions (~2–1 Ma).     

Planktonic foraminiferal turnover, diversity fluctuations and geochemical signals across the Eocene/Oligocene boundary in Tanzania

A major turnover in planktonic foraminifera occurred across the Eocene/Oligocene (E/O) boundary. New drill holes through the E/O boundary in southern Tanzania contain extremely well-preserved and diverse assemblages of planktonic foraminifera. Here we document a 1.2 million year record of assemblages, diversity and stable isotope fluctuations through this critical interval, which is often dissolved and/or recrystallised in carbonate-rich facies. The E/O boundary is marked by the abrupt extinction of all five remaining species of the family Hantkeninidae and a distinct size reduction in the genus Pseudohastigerina. The boundary is preceded over a short stratigraphic interval by the extinction of Turborotalia cerroazulensis, Turborotalia cocoaensis and Turborotalia cunialensis. Quantitative analysis of planktonic foraminiferal assemblages reveals significant changes in the abundance of certain species and the composition of the assemblages. We compare diversity fluctuations to the stable isotope record of Pseudohastigerina naguewichiensis and use multispecies stable isotope analyses to determine the life habitats of the most important species. A major shift in the evenness occurs at ~ 33.8 Ma associated with the extinction of the T cerroazulensis group suggesting acute ecological disturbance. We propose that the extinction of the T. cerroazulensis group at ~ 33.8 Ma was directly related to cooling of sea surface temperatures, while the extinction of Hantkeninidae was due to modifications in the thermal structure of the oceans and associated productivity changes. After the extinctions, renewed origination and diversification occurred, leading to a characteristic Oligocene planktonic foraminifer assemblage.     

Sea-surface temperature changes in the North Pacific during the Late Miocene

Twenty-seven radiolarian species and species groups are identified and used in this study. Their stratigraphic ranges span from Late Miocene to modern times and they appear to have distributions which have changed little over this interval of time. Census data for these 27 taxa in surface sediment samples are used to define six assemblages (Q-mode factors). The distribution of these assemblages are similar to modern water-mass distributions and can be statistically related to modern sea-surface temperatures.Census data on these same 27 taxa have been collected in the Upper Miocene intervals of six Pacific sites. These faunal data are described in terms of the modern radiolarian assemblages; and based on the multiple regression equations relating the modern assemblages to modern temperatures, estimates of Late Miocene temperatures are made. The estimated temperatures indicate that there is an overall cooling trend in the later part of the Miocene (magnetic Chron 8 to Chron 5), punctuated by several distinct cooling events. In August the low-latitude sites appear to be slightly cooler, and the mid-latitude sites warmer than in modern times. In February both the low- and mid-latitude sites appear to be warmer than modern temperatures at the same locations. The relatively warm temperatures near the base of the interval studied may be associated with the passage of warm tropical Atlantic waters into the eastern Pacific via the Isthmus of Panama. The general cooling trend seen in these records is thought to be associated with the gradual closure of these straits and the increased influence of the high southern latitudes.     

Biochronology and paleoclimatic implications of Middle Eocene to Oligocene planktic foraminiferal faunas

Planktic foraminiferal assemblages have been analyzed quantitatively in six DSDP sites in the Atlantic (Site 363), Pacific (Sites 292, 77B, 277), and Indian Ocean (Sites 219, 253) in order to determine the nature of the faunal turnover during Middle Eocene to Oligocene time. Biostratigraphic ranges of taxa and abundance distributions of dominant species are presented and illustrate striking similarities in faunal assemblages of low latitude regions in the Atlantic, Pacific and Indian oceans. A high resolution biochronology, based on dominant faunal characteristics and 55 datum events, permits correlation between all three oceans with a high degree of precision. Population studies provide a view of the global impact of the paleoclimatic and paleoceanographic changes occurring during Middle Eocene to Oligocene time.Planktic foraminiferal assemblage changes indicate a general cooling trend between Middle Eocene to Oligocene time, consistent with previously published oxygen isotope data. Major faunal changes, indicating cooling episodes, occur, however, at discrete intervals: in the Middle Eocene 44-43 Ma (P13), the Middle/Late Eocene boundary 41-40 Ma ( ), the Late Eocene 39-38 Ma ( ), the Eocene/Oligocene boundary 37-36 Ma (P18), and the Late Oligocene 31-29 Ma ( ). With the exception of the boundary, faunal changes occur abruptly during short stratigraphic intervals, and are characterized by major species extinctions and first appearances. The Eocene/Oligocene boundary cooling is marked primarily by increasing abundances of cool water species. This suggests that the boundary cooling, which marks a major event in the oxygen isotope record affected planktic faunas less than during other cooling episodes. Planktic foraminiferal faunas indicate that the boundary event is part of a continued cooling trend which began during the Middle Eocene.Two hiatus intervals are recognized in low and high latitude sections at the Middle/Late Eocene boundary and in the Late Eocene ( ). These hiatuses suggest that vigorous bottom water circulation began developing in the Middle Eocene, consistent with the onset of the faunal cooling trend, and well before the development of the psychrosphere at the boundary.     

The explosive diversification of the diatom genus Chaetoceros across the Eocene/Oligocene and Oligocene/Miocene boundaries in the Norwegian Sea

Eocene to middle Miocene stratigraphic changes in species richness, abundance and valve size of Chaetoceros resting spores in the Norwegian Sea (DSDP Site 338) were investigated in order to understand past productivity and paleoenvironmental changes in upwelling regions. As a result, drastic resting spore events were recognized in a 6 myr interval across the Eocene/Oligocene boundary (EO Event), the Oligocene/Miocene boundary (OM Event) and in the early middle Miocene (emM Event). The EO Event was characterized by explosive diversification at both the morpho-generic and specific levels, an increase in abundance, and a decrease in valve size from the upper Eocene through the lowest Oligocene. The OM Event was defined by a two-fold increase in species richness. During the emM Event spore abundance decreased rapidly, and species richness and valve size decreased gradually. These changes may indicate changes in the nutrient supply, especially in upwelling regions. The increased species richness suggests a change from a stable water column with a constant nutrient supply in the Eocene to an unstable one with a sporadic nutrient supply by increased vertical mixing in the Oligocene, based on evaluation of the ecologic differences between dinoflagellate cysts and Chaetoceros resting spores. The role of main primary producer might have switched from dinoflagellates and/or nannoplankton in the Eocene to diatoms, especially Chaetoceros, in the Oligocene in the Norwegian Sea. Increased resting spore species richness during the OM Event may show that environmental changes such as global cooling and nutrient mixing led to a diversification of the spore producing genus Chaetoceros. The emM Event might have been affected by changes in paleoceanographic conditions, perhaps a decrease in nutrient supply. This study presents the first paleoceanographic analysis using not only the total resting spore abundance but also the abundances of individual species, and establishes the value of spore taxonomy and diatom analysis including spores.     

Relationships between the spread of Caulerpa filiformis and fish communities on temperate rocky reefs

The previously sub‐dominant native marine macrophyte Caulerpa filiformis is now dominant on many sub‐tidal rocky reefs in New South Wales (NSW), Australia and is expanding its distribution. As C. filiformis is highly chemically defended and structurally different to co‐occurring habitat‐forming macrophytes, two key attributes that govern fish assemblages, we hypothesized that fish assemblages, particularly herbivorous fishes, would be different at sites where C. filiformis occurred from where it was previously absent and within sites, fish community structure would be correlated to the cover of C. filiformis. We investigated these hypotheses by determining reef‐associated fish assemblage attributes (assemblage structure, species richness, total abundance, Shannon‐Weiner diversity, abundance of herbivorous species) along transects within sites where C. filiformis was present and absent. Surprisingly, despite large patches and very high densities of C. filiformis on the reefs we sampled, at larger spatial scales (i.e., among sites) no fish assemblage metrics differed between sites with large stands of C. filiformis and sites without the alga. Moreover the abundance of one dominant herbivore, the rock cale Aplodactylus lophodon, was greater at sites within large beds of C. filiformis. At smaller spatial scales, however, i.e. within sites where C. filiformis was present, fish assemblages did vary as a function of C. filiformis cover along transects, although this was not consistent across sampling times. Overall, our results suggest that the potential effects of the spread of this alga on faunal communities warrants further investigation.     

Miocene snakes from northeastern Kazakhstan: new data on the evolution of snake assemblages in Siberia

Abstract

The Neogene snake fauna from the central and eastern regions of Eurasia is still largely unknown. This paper reports on a unique snake fauna from the late middle Miocene of the Baikadam and Malyi Kalkaman 1 and 2 localities, northeastern Kazakhstan, which represents the best-documented Miocene snake assemblage in Central Asia. Previous studies admitted that snake fauna could be homogeneous over a large part of Eurasia during the Miocene, with the late middle to early late Miocene assemblages similar to snake assemblages that inhabited Europe in the late early and early middle Miocene. This assumption is partially supported by the presence of Texasophis bohemiacus and Coluber cf. hungaricus, as well as vipers of the ‘V. aspis’ complex. However, the presence of taxa which are (1) probably not related to European representatives (‘Colubrinae’ A and B), (2) probably never occurred in Central and Western Europe and (3) are closely related to species recently inhabiting southern Siberia (Elaphe aff. dione, Gloydius sp.) indicates that faunal dissimilarity was relatively high within Eurasia during the late middle Miocene. This assumption is in accordance with studies of small mammal assemblages which show a decreasing homogenity in the Eurasia in the course of the middle Miocene.     

Vertical Distribution of Radiolarians and Their Role in Epipelagic Communities of the East Pacific Rise and the Gulf of California

Vertical distribution of different groups of radiolarians and mesozooplankton (l = 0.2–3.0 mm) and the contribution of radiolarians to total biomass of bathometer-sampled zooplankton were studied in the epipelagic East Pacific Rise and the Gulf of California during 49th cruise of R/V “Akademik Mstislav Keldysh” (September–October, 2003). The production of the symbiotic algae of acantharians and colonial radiolarians as well as their contribution to the total primary production were evaluated. The proportion of all radiolarian groups in the total zooplankton biomass was shown to range from 1.4 to 11.5%. The contribution of zooxanthellae to the total primary production was not great in the studied region and equaled 0.1–1.2 and 0.3–0.7% for colonial radiolarians and acantharians, respectively.__________Translated from Izvestiya Akademii Nauk, Seriya Biologicheskaya, No. 3, 2005, pp. 339–348.Original Russian Text Copyright © 2005 by Zasko, Rusanov.