Phenotypic evolution in a fossil gastropod species lineage: evidence for adaptive radiation?

Detecting speciation in the fossil record is a particularly challenging matter. Palaeontologists are usually confronted with poor preservation and limited knowledge on the palaeoenvironment. Even in the contrary case of adequate preservation and information, the linkage of pattern to process is often obscured by insufficient temporal resolution. Consequently, reliable documentations of speciation in fossils with discussions on underlying mechanisms are rare. Here we present a well-resolved pattern of morphological evolution in a fossil species lineage of the gastropod Melanopsis in the long-lived Lake Pannon. These developments are related to environmental changes, documented by isotope and stratigraphical data. After a long period of stasis, the ancestral species experiences a phenotypic change expressed as shift and expansion of the morphospace. The appearance of several new phenotypes along with changes in the environment is interpreted as adaptive radiation. Lake-level high stands affect distribution and availability of habitats and, as a result of varied functional demands on shell geometry, the distribution of phenotypes. The on-going divergence of the morphospace into two branches argues for increasing reproductive isolation, consistent with the model of ecological speciation. In the latest phase, however, progressively unstable conditions restrict availability of niches, allowing survival of one branch only.

Pleistocene sedimentation in the Benguela Current, southeastern Atlantic

We infer variations in paleoproductivity and eolian input at ODP Site 1082 in the Walvis Basin from stable oxygen isotope compositions of the planktonic foraminifera Globorotalia inflata, total organic carbon mass accumulation rates (TOC MAR), and X-ray fluorescence analyses of Fe content. The most pronounced paleoclimatic changes correspond to the time at about 0.9 Ma, when glacial conditions in the northern hemisphere (NH) led to the onset of pronounced 100-kyr glacial-interglacial cycles. We used Fe intensity as a proxy for eolian terrigenous input, and TOC MAR as a paleoproductivity indicator. Paleoproductivity and eolian input show generally higher-amplitude variations of glacial-interglacial cyclicity from 1.5 to 0.58 Ma, indicating pronounced variations in upwellingfavorable winds in this area. At 0.58 Ma, paleoproductivity and eolian input shifted abruptly to lower-amplitude variations with a periodicity of 100 kyr while delta18O values show a trend toward more negative isotope values for the past 0.65 Myr. Especially during glacial periods, oxygen isotope values indicate increasingly warmer sea-surface temperatures toward the end of the Pleistocene. To evaluate the relative influences of NH glaciation and southern hemisphere (SH) insolation as potential forcing mechanisms for variations of eolian input and productivity in the northern Benguela system, we filtered our proxy records at orbital frequencies. The filtered records of Fe intensity and TOC MAR indicate a strong influence of the 100-kyr and 41-kyr frequency bands, supporting our assumption that strong ice buildup in the NH is the dominant trigger for climate changes on the continent and probably in trade-wind intensity. SH insolation and low-latitude precession-related insolation changes were important for paleoproductivity variations in the northern Benguela system, modifying the nutrient supply by southern ocean intermediate waters and the zonal direction of upwelling-inducing trades by the African monsoon system, respectively.

Distribution of diatoms in Oligocene to Pleistocene sediments of the tropical Pacific

According to the drilling probes of the Deep Waier Drilling Project, Neogene sediments in a tropical area of the Pacific Ocean are divided into 15 zones based on diatoms. The author shows that a unique zonation may be applied for the entire region. Identification of diatoms zones boundaries was conducted through their direct correlation with nannoplancton, radiolarian and foraminiferal zonal sceals. Their ultra-structure and morphological relationship are being analysed. The mode of siliceous accumulation within the equatorial belt differed through the western central and eastern region since the early Miocene and the difference become more evident from the end of Middle Miocene. The distribution of Neogene diatomaceous silt in the tropical area is controlled by the character of gyre-water circulation and agrees with the modern geographical zonation.

Plio-Pleistocene magnetic polarity stratigraphies of ODP Leg 177 Sites from the South Atlantic

Magnetic polarity stratigraphies from ODP Leg 177 'high resolution' sites indicate Brunhes sedimentation rates in the 12-25 cm/kyr range, with a trend of decreasing sedimentation rates with increasing age. Magnetite is the principal remanence-carrying mineral. Downcore alteration of magnetite and authigenic growth of iron sulfides introduces a high coercivity diagenetic remanence carrier (pyrrhotite). The change in pore water sulfate with depth in the sediment tends to be in step with the decrease in magnetization intensity, indicating the link between sulfate reduction and magnetite dissolution. Shipboard pass-through magnetometer data are generally very noisy due to a combination of weak magnetization intensities, drilling-related core deformation, and the influence of authigenic iron sulfides. Post-cruise progressive demagnetization of discrete samples aids the magnetostratigraphic interpretation, as these measurements are less influenced by low magnetization intensities and drilling-related deformation. The magnetostratigraphic interpretations provide much-needed calibration for biostratigraphic events in the high latitude southern oceans. Apart from the ODP Hole 745B (Kerguelen Plateau), published Plio-Pleistocene magnetostratigraphies from ODP sites in the Southern Ocean are poorly constrained. For this reason, we compare interpolated ages of 11 radiolarian events and one diatom event that occur at Hole 745B and Leg 177 sites.

Diatom abundance in middle-to-late Pleistocene sediments of IODP Site 303-U1304

We examined diatom assemblages in a series of remarkable laminated diatomaceous ooze (LDO) horizons in the marine sediments from Integrated Ocean Drilling Program (IODP) Site U1304 to reconstruct the middle-to-late Pleistocene paleoceanographic evolution of the northern North Atlantic Ocean. Four confirmed diatom biohorizons combined with calcareous nannofossil and paleomagnetic stratigraphies established the chronological framework for the material. The planktonic, araphid, needle-like species Thalassiothrix longissima was the greatest contributor to the LDO facies. From the results of a principal component analysis using the percent abundances of 65 significant (p = 5%) diatom taxa, except for Tx. longissima, which was extremely dominant in almost all horizons observed, we identified two principal component (PC) axes. Taxa probably associated with the stratigraphic distribution of the major zonal marker Neodenticula seminae (ranging from 1.26 to 0.84 Ma in this ocean) loaded on PC1 with a high value. PC2 was related to the ocean surface temperature. The stratigraphic variability of the PC2 score indicated that switching between warm- and cold-water assemblages occurred concurrently with LDO deposition (or extreme Tx. longissima dominance) episodes in several horizons (particularly after 0.84 Ma), suggesting that the Subarctic Convergence (SAC) oceanic front passed over Site U1304 during Pleistocene glacial/interglacial cycles. Our floral evidence supports the model of nearly monospecific LDO formation caused by the enhanced physical accumulation of particular diatoms such as Tx. longissima. On the other hand, Nd. seminae, which probably contributes to spring phytoplankton blooms in the modern ocean, was present only between 1.26 and 0.84 Ma in this area. Thus, we infer that the main contributor of export flux in the regional annual primary production cycle would have shifted drastically from one of a spring phytoplankton bloom leader (Nd. seminae) to minor but mass dump assemblages (Tx. longissima etc.) in the mid-Pleistocene.

Distribution of Cycladophora davisiana davisiana in upper Pliocene and Pleistocene sediments of the North Atlantic and Labrador Sea

Upper Pliocene and Pleistocene abundance fluctuations of the radiolarian Cycladophora davisiana (Ehrenberg) davisiana (Petrushevskaya) are documented from North Atlantic (Site 609) and Labrador Sea (Site 646B) to provide the first long-term correlation of its abundance fluctuations to oxygen isotope stages 1-114. Also examined are temporal and regional fluctuations in abundances C. d. davisiana and the global dispersal routes of the species. The first occurrence of C. d. davisiana in the eastern North Atlantic Ocean (Site 609) occurred between 2.586 and 2.435 Ma (oxygen isotope stages 109.66-102.19). During the early Matuyama Chron, prior to oxygen isotope stage 63, C. d. davisiana abundances were less than 1% and never greater than 12%, while abundances of greater than 5% are found in stages 65.71-73, 74, and 83-84. The initial major abundance peak (35.7%) of C. d. davisiana was noted near the stage 63/62 boundary. Abundance peaks of greater than 15%, between oxygen isotope stages 35 and 63, are limited to stages 63.02, 58.07, 55.07-54.26, and 50.76-50.22. These represent the only such abundance peaks detected during the first c. 1.5 million years of the species within the North Atlantic. The character of C. d. davisiana abundance fluctuations in Site 609 changes after oxygen isotope stage 35; average abundances are greater (7.7% vs. 4.3%) and abundance maxima of more than 15% are more frequent. Many, but not all, peak abundances of C. d. davisiana occur in glacial stages (e.g., 8, 14, 18, 20, 26, 30, 34, 50, 54, and 58). Increased abundances of the species are also noted in weak interglacial stages (e.g., stages 3, 23, 39, and 41), and significant cool periods of robust interglacial periods (e.g., late stage 11). Sample spacing is adequate in some stages to note some rapid changes in abundance near stage transitions (e.g., stages 4/5, 25/26, 62/63). The sample density in Holes 609 and 611 and the upper portion of 646B is sufficient to detect a synchroneity of many abundance maxima and minima among sites. Some abundance peaks are undetected in one or more of the two holes, warranting further sampling to obtain a more accurate record of regional abundance fluctuations. Prior to stage 36, few ages of Hole 611 peaks are the same as those in the more precisely dated Hole 609. The highest abundances of C. d. davisiana were noted in Labrador Sea Hole 646B where the earliest known occurrence of the species is documented (3.08-2.99 Ma). C. d. davisiana is inferred to have evolved in the Labrador Sea (or Arctic), and migrated next through the Arctic into the North Pacific (2.62-2.64 Ma, stage 114) before migrating into the Norwegian Sea (2.63-2.53 Ma) and North Atlantic (2.59-2.44 Ma, stages 109-102). Additional migration of C. d. dauisiana into the southern South Atlantic (Site 704) occurred much later (2.06 Ma, stage 83).

Revision of the early-middle Pleistocene calcareous nannofossil biochronology

The extant nannofossil biostratigraphic and biochronologic framework for the early-middle Pleistocene time interval has been tested through the micropaleontological analysis of globally distributed high-quality low- to mid-latitude deep-sea successions. The quantitative temporal distribution patterns of relative abundances of selected taxa were reconstructed in critical intervals, and the following biohorizons were defined: first occurrence of medium-sized Gephyrocapsa spp. (bmG); last occurrence of Calcidiscus macintyrei (tCm); first occurrence of large Gephyrocapsa spp. (blG); last occurrence of large Gephyrocapsa spp. (tlG); first occurrence of Reticulofenestra asanoi (bRa); re-entrance of medium-sized Gephyrocapsa spp. (reemG) and last occurrence of Reticulofenestra asanoi (tRa). The detailed patterns of abundance change at these biohorizons were used to generate a detailed biostratigraphy, and the biostratigraphic data were transformed into a precise biochronology by means of correlation to isotope stratigraphies and astronomical timescales. The degree of isochrony or diachrony of the biohorizons was evaluated. Biohorizons tlG and tRa are isochronous occurring close to marine isotope stages (MIS)55 and MIS 22, respectively, and bmG and blG are slightly diachronous on the order of 30-40 kyr, whereas biohorizons tCm, reemG and bRa are confirmed as diachronous on the order of 100, 80 and 60 kyr, respectively. Some of the events are clearly controlled by environmental conditions, e.g. the last occurrence of R. asanoi, related to significant environmental changes associated with the first large-amplitude glaciation of the late Quaternary, MIS 22.