The evolutionary history of size variation of planktic foraminiferal assemblages in the Cenozoic

The iterative evolutionary radiation of planktic foraminifers is a well-documented macroevolutionary process. Here we document the accompanying size changes in entire planktic foraminiferal assemblages for the past 70 My and their relationship to paleoenvironmental changes. After the size decrease at the Cretaceous/Paleogene (K/P) boundary, high latitude assemblages remained consistently small. Size evolution in low latitudes can be divided into three major phases: the first is characterized by dwarfs (65-42 Ma), the second shows moderate size fluctuations (42-14 Ma), and in the third phase, planktic foraminifers have grown to the unprecedented sizes observed today. Our analyses of size variability with paleoproxy records indicate that periods of size increase coincided with phases of global cooling (Eocene and Neogene). These periods were characterized by enhanced latitudinal and vertical temperature gradients in the oceans and high diversity (polytaxy). In the Paleocene and during the Oligocene, the observed (minor) size changes of the largely low-diversity (oligotaxic) assemblages seem to correlate with productivity changes. However, polytaxy per se was not responsible for larger test sizes.

Ocean acidification and its potential effects on the early life-history of non-calcifying and calcifying echinoderm (Odontaster validus, Patiriella regularis and Arachnoides placenta) larvae, 2011

Ocean acidification, as a result of increased atmospheric CO2, has the potential to adversely affect the larval stages of many marine organisms and hence have profound effects on marine ecosystems. This is the first study of its kind to investigate the effects of ocean acidification on the early life-history stages of three echinoderms species, two asteroids and one irregular echinoid. Potential latitudinal variations on the effects of ocean acidification were also investigated by selecting a polar species (Odontaster validus), a temperate species (Patiriella regularis), and a tropical species (Arachnoides placenta). The effects of reduced seawater pH levels on the fertilization of gametes, larval survival and morphometrics on the aforementioned species were evaluated under experimental conditions. The pH levels considered for this research include ambient seawater (pH 8.1 or pH 8.2), levels predicted for 2100 (pH 7.7 and pH 7.6) and the extreme pH of 7.0, adjusted by bubbling CO2 gas into filtered seawater. Fertilization for Odontaster validus and Patiriella regularis for the predicted scenarios for 2100 was robust, whereas fertilization was significantly reduced in Arachnoides placenta. Larval survival was robust for the three species at pH 7.8, but numbers declined when pH dropped below 7.6. Normal A. placenta larvae developed in pH 7.8, whereas smaller larvae were observed for O. validus and P. regularis under the same pH treatment. Seawater pH levels below 7.6 resulted in smaller and underdeveloped larvae for all three species. The greatest effects were expected for the Antarctic asteroid O. validus but overall the tropical sand dollar A. placenta was the most affected by the reduction in seawater pH. The effects of ocean acidification on the asteroids O. validus and P. regulars, and the sand dollar A. placenta are species-specific. Several parameters, such as taxonomic differences, physiology, genetic makeup and the population's evolutionary history may have contributed to this variability. This study highlights the vulnerability of the early developmental stages and the complexity of ocean acidification. However, future research is needed to understand the effects at individual, community and ecosystem levels.

Changes in calcareous nannofossil assemblages across the Paleocene/Eocene transition from the paleo-equatorial Pacific Ocean

Quantitative analyses of selected calcareous nannofossils in deep-sea sections recovered from the paleo-equatorial Pacific (ODP Leg 199) provide new information about biostratigraphy, biochronology and the evolutionary history of calcareous nannofossils across the Paleocene/Eocene transition interval. The sediment cores from ODP Leg 199 represent the first continuous Paleocene/Eocene boundary sections ever to be sampled in the central equatorial Pacific Ocean. Calcareous nannofossil assemblages are studied to document the distribution of biostratigraphically useful taxa such as Ericsonia, Discoaster, Fasciculithus, Rhomboaster and Tribrachiatus. Focus is given to the evolution of the Rhomboaster-Tribrachiatus lineage in the lower Eocene interval at Site 1215, and on the stratigraphic relationship of these taxa relative to species in the genus Fasciculithus. Critical intervals of North Atlantic DSDP Site 550 have also been re-examined. The Tribrachiatus digitalis morphotype was described at Site 550 from an interval affected by down-hole contamination, partly originating from within the Tribrachiatus orthostylus range. The T. digitalis morphotype represents an evolutionary transitional form between T. contortus and T. orthostylus, entering the stratigraphic record within the range of the former species and disappearing within the lower part of the range of the latter species. The subzonal subdivision of Zone NP10 hence collapses. Lithological and colour variability reflecting orbital cyclicity occur in the lower Eocene of Site 1215, permitting a relative astronomical age calibration of the Tribrachiatus taxa. The distinct Rhomboaster spp.-Discoaster araneus association also occurs in the paleo-equatorial Pacific Ocean, together with a marked decrease in diversity of Fasciculithus spp. Site 1220 reveals a short peak abundance of Thoracosphaera spp. just above the P/E boundary interval, which probably reflects a stressed surface water environment.

Contemporary genetic, historical genetic and morphological data sets for the Yellow-tufted Honeyeater Lichenostomus melanops

Understanding the evolutionary history of threatened populations can improve their conservation management. Re-establishment of past but recent gene flow could re-invigorate threatened populations and replenish genetic diversity, necessary for population persistence. One of the four nominal subspecies of the common yellow-tufted honeyeater, Lichenostomus melanops cassidix, is critically endangered despite substantial conservation efforts over 55 years. Using a combination of morphometric, genetic and modelling approaches we tested for its evolutionary distinctiveness and conservation merit. We confirmed that cassidix has at least one morphometric distinction. It also differs genetically from the other subspecies in allele frequencies but not phylogenetically, implying that its evolution was recent. Modelling historical distribution supported the lack of vicariance and suggested a possibility of gene flow among subspecies at least since the late Pleistocene. Multi-locus coalescent analyses indicated that cassidix diverged from its common ancestor with neighbouring subspecies gippslandicus sometime from the mid-Pleistocene to the Holocene, and that it has the smallest historical effective population size of all subspecies. It appears that cassidix diverged from its ancestor with gippslandicus through a combination of drift and local selection. From patterns of genetic subdivision on two spatial scales and morphological variation we concluded that cassidix, gippslandicus and (melanops + meltoni) are diagnosable as subspecies. Low genetic diversity and effective population size of cassidix may translate to low genetic fitness and evolutionary potential, thus managed gene flow from gippslandicus is recommended for its recovery.

Origin and evolution of Cycladophora davisiana (Radiolaria) in DSDP Hole 19-192, Northwest Pacific

This paper documents the evolutionary history of Cycladophora davisiana Ehrenberg from an uppermost Miocene to Pleistocene sedimentary record in the high-latitude Northwest Pacific. It apparently evolved from C. sakaii Motoyama through a series of intermediates. C. sakaii has a relatively large shell with an external spongy layer. The evolutionary transition is characterized by a relatively rapid decrease in thorax size with a reduction of the spongy appendage. This change occurred during about 0.4 m.y. from 2.8 to 2.4 Ma without cladogenesis. Following this interval, a decrease in thorax size continued gradually up to the Recent, resulting in a very small morphology. Although the population of C. davisiana first appeared at about 2.5 Ma, some morphotypic specimens may occur in earlier periods as indistinguishable very small endmembers in the C. sakaii populations. Timing of the first appearance events both of morphotypic specimens and of a population of C. davisiana in Site 192 and previously reported cores does not disprove the idea that C. davisiana evolved first in the Northwest Pacific region, and later migrated into other regions of the world ocean. Biometrics clearly indicate no direct phylogenetic relationships between C. davisiana and C. cornutoides Kling in the studied core. Thus, the latter species, which was originally described as a variation and later elevated to a subspecies of the former species, is separated from the former species and raised to the species rank.

Petrology and carbon and oxygen stable isotopic composition of macrofossils and sediments from the Blake-Bahama Formation, DSDP Hole 93-603B

Strata that record the evolutionary history of the North American continental margin in a region that serves as the basin margin interface between allochthonous sedimentation from the continent and pelagic sedimentation from the oceanic realm were recovered at Deep Sea Drilling Project Site 603, on the lower continental rise. The lowermost unit recovered at this site is composed of upper Berriasian-Aptian interbedded laminated limestone and bioturbated limestone with sandstone to claystone turbidites. This unit can be correlated with the Blake-Bahama Formation in the western North Atlantic. Studies of the laminated and bioturbated limestones were used to determine the depositional environment. Geochemical and petrographic studies suggest that the laminated limestones were deposited from the suspended particulate loads of the nepheloid layer associated with weak bottom-current activity as well as moderate to poorly oxygenated bottom-water conditions. Fragments of macrofossils are also found in the Blake-Bahama Formation drilled at Site 603. Twelve specimens and their host sediment were analyzed for their carbon and oxygen isotopic composition. The macrofossil samples chosen for analysis consist of nine samples of Inoceramus, two ammonite aptychi, and one belemnite sample. Depletion in 18O is observed in recrystallized specimens. The ammonite aptychi have been diagenetically altered and/or exhibit evidence of isotopic fractionation by the organism. Oxygen isotope paleotemperatures obtained from five well-preserved specimens - four of Inoceramus and one of a belemnite - suggest that bottom-water temperatures in the North Atlantic Basin during the Early Cretaceous were very warm, at least 11°C.

Planktonic foraminiferal stratigraphy and datum levels of ODP Leg 182 sites

Planktonic foraminifers from Ocean Drilling Program Leg 182, Holes 1126B and 1126C, 1128B and 1128C, 1130A and 1130B, 1132B, and 1134A and 1134B confirm the neritic record that during the early Miocene the Great Australian Bight region was in a cool-temperate regime with abundant Globoturborotalita woodi. Warm marine environments started to develop in the later part of the early Miocene, and the region became warm temperate to subtropical in the early middle Miocene with abundant Globigerinoides, Orbulina, and Globorotalia, corresponding to global warming at the Miocene climatic optimum. Fluctuations between cool- and warm-temperate conditions prevailed during the late Miocene, as indicated by abundant Globoconella conoidea and Menardella spp. A major change in planktonic foraminiferal assemblages close to the Miocene/Pliocene boundary not only drove many Miocene species into extinction but also brought about such new species as Globorotalia crassaformis and Globoconella puncticulata. Warm-temperate environments continued into the early and mid-Pliocene before being replaced by cooler conditions, supporting numerous Globoconella inflata and Globigerina quinqueloba. Based on data from this study and published results from the Australia-New Zealand region, we established a local planktonic foraminifer zonation scheme for separating the southern Australian Neogene (SAN) into Zones SAN1 to SAN19 characterizing the Miocene and Zones SAN20 to SAN25 characterizing the Pliocene. The Neogene sections from the Great Australian Bight are bounded by hiatuses of ~0.5 to >3 m.y. in duration, although poor core recovery in some holes obscured a proper biostratigraphic resolution. A total of 15 hiatuses, numbered 1 to 15, were identified as synchronous events from the base of the Miocene to the lower part of the Pleistocene. We believe that these are local manifestations of major third-order boundaries at about (1) 23.8, (2) 22.3, (3) 20.5, (4) 18.7, (5) 16.4, (6) 14.8, (7) 13.5, (8) 11.5, (9) 9.3, (10) 7.0, (11) 6.0, (12) 4.5, (13) 3.5, (14) 2.5, and (15) 1.5 Ma, respectively. This hiatus-bounded Neogene succession samples regional transgressions and stages of southern Australia and reveals its stepwise evolutionary history.

(Table 1) Number of alleles per locus and heterozygosity of Arctic charr (Salvelinus alpinus) populations in Iceland

The ecological theory of adaptive radiation predicts that the evolution of phenotypic diversity within species is generated by divergent natural selection arising from different environments and competition between species. Genetic connectivity among populations is likely also to have an important role in both the origin and maintenance of adaptive genetic diversity. Our goal was to evaluate the potential roles of genetic connectivity and natural selection in the maintenance of adaptive phenotypic differences among morphs of Arctic charr, Salvelinus alpinus, in Iceland. At a large spatial scale, we tested the predictive power of geographic structure and phenotypic variation for patterns of neutral genetic variation among populations throughout Iceland. At a smaller scale, we evaluated the genetic differentiation between two morphs in Lake Thingvallavatn relative to historically explicit, coalescent-based null models of the evolutionary history of these lineages. At the large spatial scale, populations are highly differentiated, but weakly structured, both geographically and with respect to patterns of phenotypic variation. At the intralacustrine scale, we observe modest genetic differentiation between two morphs, but this level of differentiation is nonetheless consistent with strong reproductive isolation throughout the Holocene. Rather than a result of the homogenizing effect of gene flow in a system at migration-drift equilibrium, the modest level of genetic differentiation could equally be a result of slow neutral divergence by drift in large populations. We conclude that contemporary and recent patterns of restricted gene flow have been highly conducive to the evolution and maintenance of adaptive genetic variation in Icelandic Arctic charr.

Late Oligocene rapid transformations in the South China Sea

Lithobiostratigraphic data indicate that the double reflectors on the seismic profile through Ocean Drilling Program (ODP) Site 1148 represent two unconformities that coincide, respectively, with the lower/upper Oligocene boundary at ~488 mcd, and Oligocene-Miocene boundary at 460 mcd. Two other unconformities, at ~478 and 472 mcd, respectively, were also identified within the upper Oligocene section. Together they erased a sediment record of about 3 Ma from this locality in a period of very active seafloor spreading. The existence of 32.8 Ma marine sediment at the terminated depth (850 mcd) indicates that the initial breakup of the South China Sea (SCS) was probably during 34-33 Ma, close to the Eocene-Oligocene boundary. High sedimentation rates of 60-115 m/my from the much expanded, N350 m lower Oligocene section resulted from rifting and rapid subsidence between 33 and 29 Ma. The mid-Oligocene unconformity at ~28.5 Ma, which also occurred in many parts of the Indo-West Pacific region, was probably related to a significant uplift of the Himalayan-Tibetan Plateau to the west and the initial collision between Indonesia and Australia in the south. A narrowed Indonesian seaway may have accounted for the late Oligocene warming and chalk deposition in the northern South China Sea including the Site 1148 locality. The unconformities and slumps near the Oligocene-Miocene boundary indicate a very unstable tectonic regime, probably corresponding to changes in the rotation of different land blocks and the seafloor spreading ridge from nearly E-W to NE-SW, as recognized earlier at magnetic Anomaly 7. This 25 Ma event also saw the first New Guinea terrane docking at the northern Australian craton. The low sedimentation rate of ~15 m/my in the early to middle Miocene may correspond to another period of rapid seafloor spreading and rapid widespread subsidence that effectively caused sediment source areas to retreat with a rapidly rising sea level. The isostatic nature of these late Oligocene unconformities and slumps with several major collision-uplift events indicate that the rapid changes in the early evolutionary history of the South China Sea were mainly responding to regional tectonic reconfiguration including the uplift-driven southeast extrusion of the Indochina subcontinent.