Glacial oceanographic contrasts explain phylogeography of Australian bull kelp

The evolutionary effects of Southern Hemisphere Pleistocene oceanographic conditions — marked by fluctuations in sea levels and water temperatures, and redirected currents — are poorly understood. The southeastern tip of Australia presents an intriguing model system for studying the biological impacts of palaeoceanography. In particular, contrasting oceanographic conditions that existed on eastern vs. western sides of the Bassian Isthmus during Pleistocene glacial periods allow for natural comparisons between putative refugial vs. re-invading populations. Whereas many western Tasmanian marine taxa were likely eliminated by cold subantarctic water during the last glacial period, eastern Tasmanian populations would have persisted in relatively warm temperatures mediated by the ongoing influence of the East Australian Current (EAC). Here we test for the effects of contrasting palaeoceanographic conditions on endemic bull kelp, Durvillaea potatorum , using DNA sequence analysis (COI; rbc L) of more than 100 individuals from 14 localities in southeastern Australia. Phylogenetic reconstructions reveal a deep (maximum divergence 4.7%) genetic split within D. potatorum , corresponding to the 'eastern' and 'western' geographical regions delimited by the Bassian Isthmus, a vicariant barrier during low Pleistocene sea levels. Concordant with the western region's cold glacial conditions, samples from western Tasmania and western Victoria are genetically monomorphic, suggesting postglacial expansion from a mainland refugium. Eastern samples, in contrast, comprise distinct regional haplogroups, suggesting the species persisted in eastern Tasmania throughout recent glacial periods. The deep east–west divergence seems consistent with earlier reports of morphological differences between 'western' and 'eastern' D. potatorum , and it seems likely that these forms represent reproductively isolated species.     

Mesozoic calcareous nannofossils — state of the art

Calcareous nannofossils originated in the Triassic, radiated in the Jurassic and became a dominant component of the marine biosphere from the earliest Jurassic onward. They can be considered as one of the most important “innovations” of the Mesozoic oceans. Their basic morphology allows the differentiation of three different groups: coccoliths, nannoliths and calcispheres (= calcareous dinocysts). Only coccoliths and nannoliths are discussed in this article in some detail. Coccoliths and nannoliths have contributed greatly in the Interpretation of Mesozoic marine Systems through biostratigraphy and palaeoecology/palaeoceanography. Ever since the late 1960s both coccoliths and nannoliths have proven to be useful and reliable zonal markers for biostratigraphic schemes, allowing detailed zonations for the Jurassic and Cretaceous. Though affected by palaeobiogeographic provincialism, coccoliths and nannoliths have supplied many cosmopolitan biostratigraphic markers. These allow a global correlation of marine sedimentary units both from onshore sections in the classical European and North American areas and pelagic sequences recovered in the course of the DSDP/ODP drilling from the world’s oceans. Thus research on calcareous nannofossils Covers both, regional and global aspects. Research in the last 15 years concentrated on palaeoecological aspects. Apart from dinoflagellates, coccolithophores were the most important primary producers in Mesozoic oceans. As such they heavily relied on autecological factors such as light, nutrients and temperature. Variations in the assemblage composition of these groups may thus be viewed as a key for understanding palaeoecological, palaeoceanographic and palaeoclimatic changes of the past.      

Evidence of increasing surface water oligotrophy during the Campanian–Maastrichtian boundary interval: Calcareous nannofossils from DSDP Hole 390A (Blake Nose)

The latest Cretaceous (Campanian–Maastrichtian) is characterized by several global cooling and intermittent warming events. These climatic changes influenced the palaeoceanography substantially, including changes of the deep water sources and surface water currents. One of the most prominent episodes of climatic cooling occurred during the Campanian–Maastrichtian transition. This study focuses on the palaeoclimate and palaeoceanography of the Campanian–Maastrichtian transition by analysing the calcareous nannofossils of DSDP Hole 390A (139.92–126.15 mbsf; Blake Nose). For the examination of calcareous nannofossils sixty samples were processed using the settling technique. Biostratigraphical index taxa (Broinsonia parca constricta, Uniplanarius trifidus, and Tranolithus orionatus) suggest a late Campanian age for the major part of the studied section. The calcareous nannofossils are well preserved, highly abundant (6.80 billion specimens/gram sediment) and diverse (80 species/sample). The assemblages are dominated by Prediscosphaera spp. (20.5%), Watznaueria spp. (20.3%) and Retecapsa spp. (9.8%). Cool water taxa (Ahmuellerella octoradiata, Gartnerago segmentatum, and Kamptnerius magnificus), however, appear less frequently and do not exceed more than 1%. Due to their rarity these cool water taxa do not support the existence of an intense cooling phase during the Campanian–Maastrichtian transition at DSDP Hole 390A. Around 133 mbsf several nannofossil taxa, however, show a distinctive turnover. Mesotrophic species like Discorhabdus ignotus, Zeugrhabdotus bicrescenticus and Zygodiscus exmouthiensis are abundant below 133 mbsf, whereas oligotrophic taxa like Watznaueria spp., Eiffellithus spp. and Staurolithites flavus become common above this level. These changes imply a decrease in the input of nutrients, perhaps caused by a reorganization of ocean currents (Palaeo Gulf Stream) and reduced upwelling.     

Protoperidiniacean dinoflagellate cyst taxa from the Upper Miocene of ODP Leg 178, Antarctic Peninsula

Protoperidiniacean dinoflagellate cysts have been recovered from Upper Miocene sediments of Hole 1095, ODP Leg 178, drilled to the west of the Antarctic Peninsula. These cysts make up virtually the entire dinoflagellate cyst assemblages, and three new species are formally described herein as Selenopemphix bothrion sp. nov., Selenopemphix kepion sp. nov. and Selenopemphix minys sp. nov., together with one informal taxon and two species attributed to published taxa. The occurrence of Late Miocene protoperidiniacean-dominated assemblages in the vicinity of the Antarctic continent is of special interest. Their presence may indicate the initiation of the Antarctic Circumpolar Current and increased nutrient supply within the Southern Ocean as the Antarctic Divergence became established. A comparison with several other protoperidiniacean dinoflagellate assemblages of similar age in other parts of the world provides some preliminary evidence for possible concomitant oceanographic change at this time.     

Multivariate hierarchical analyses of Early Jurassic Ostracoda assemblages

Palaeobiogeographic patterns of Early Jurassic ostracods from the northern and southern hemispheres (96 sections located in Europe, North Africa, Western Australia and North and South America) based on 243 species-level records document global patterns of distribution that can be compared to those previously published on ostracods from the European Epicontinental Sea and Tethyan and South Panthalassa areas. All described records of ostracods from both hemispheres spanning the Hettangian to Early Toarcian have been compiled and verified, and their patterns of origin and distribution have been interpreted. Jaccard coefficient of similarity was used to asses similarities among European, American and Tethyan ostracod shelf faunas. The numerical analysis shows a progressive longitudinal gradient in provincialism through the Early Jurassic, consistent with the northward drift of Tethyan ostracod faunas towards the European Epicontinental Sea and the southward movement of European taxa into Tethys and Panthalassa oceans. The spread of cosmopolitan species and extinction of endemic species, allied to the disappearance of geographical barriers, warmer climate conditions and rising sea levels can explain the reduction in ostracod diversity and the east-west provincialism throughout the Early Jurassic. Interchange between hemispheres, including bipolar distributions, are recognized from the Sinemurian time, pointing out that for most of the studied period, the climate worldwide was warm and tropical.     

Biotic signals from nannoflora across the iridium anomaly in the upper Eocene of the Massignano section: evidence from statistical analysis

Strong evidence for a major late Eocene impact, including an iridium-rich layer and shocked quartz, has been found in the Massignano section (central Italy), stratotype of the Eocene/Oligocene boundary. Quantitative calcareous nannofossils analyses have been performed and classical and multivariate statistical techniques have been applied to assess the paleonvironmental effect of this impact on the nannoflora.Abundance patterns indicate: (1) a warming episode just above the iridium layer; (2) a subsequent slight cooling trend, probably related to the global cooling during late Eocene to early Oligocene; (3) marked fluctuations in the trophic resources and an increase in near-shore taxa in the upper part of the interval; (4) increase in low-nutrient taxa in biotite-rich layers, just above the iridium level, possibly related to lowered productivity linked to volcanic activity. All these signals appear most likely to be the result of long-term climatic changes rather than to short-term effects of extraterrestrial impact.In addition, cluster analysis was performed on the assemblage data. This delineated similarities among different taxa and grouped samples with similar assemblages. Several groups related to temperature-, trophic- and unknown-preference conditions have been distinguished. A consistent difference in the abundances of the taxa and in their correlation has been detected between samples above and below the iridium level. This change appears to be related to two important sedimentary features which are found in the interval above the iridium occurrence: (1) the presence of biotite-rich layers linked to increasing volcanism and (2) a conspicuous color transition from reddish to greenish related to changes in sedimentary environments (e.g. variations in trace element abundance). These environmental changes rather than the impact may have affected the nannoflora assemblages.     

Subtropical Front fluctuations south of Australia (45°09′S, 146°17′E) for the last 130 ka years based on calcareous nannoplankton

Calcareous nannoplankton assemblages from a Late Quaternary deep-sea core (GC07; 46°09′S, 146°17′E) south of Australia provide information on regional palaeoceanography and palaeoclimate changes in the Southern Ocean, in particular the movement of the Subtropical Front for the past 130 ka years. Marine Isotope Stages 1–5 are identified through changes in calcareous nannoplankton assemblages, supported by ¹⁴C dates, and oxygen isotope and %CaCO data.Two distinct assemblages are recognised: a warm water assemblage with higher abundances of Calcidiscus leptoporus, Emiliania huxleyi, Helicosphaera.carteri, Syracosphaera pulchra, Gephyrocapsa caribbeanica and Gephyrocapsa oceanica; and, a cold water assemblage with higher abundances of Gephyrocapsa muellerae and Coccolithus pelagicus. Alternation between these two assemblages downcore in GC07 reflect movement of the Subtropical Front across the location and can be correlated to Marine Isotope Stages (MIS) 1–5. Sediments with a cold water assemblage indicate the position of the Subtropical Front equatorward of the site when transitional to sub-antarctic waters were overlying the site. Conversely sediments with a warm water assemblage indicate the Subtropical Front was poleward of GC07 when warmer, subtropical waters were over the site. MIS 1 and 5 are interpreted as warmer than MIS 3 (based on species composition) with the Subtropical Front more poleward than for MIS 3. During MIS 3 the Subtropical Front is interpreted as adjacent to or immediately poleward of GC07. Some species including C. leptoporus and C. pelagicus show negative covariance and are considered to be reliable species in identifying glacial and interglacial intervals in this region.Comparison with established biostratigraphy based on calcareous nannoplankton showed the datum event for the reversal between E. huxleyi and G. muellerae of 73 ka in transitional waters is not applicable in this region. The reversal between these two species occurs between 48 and 30 cm downcore in GC07 with a ¹⁴C date of 11 020 year BP at 49–48 cm, i.e. the reversal event is younger than this date.     

Absence of post-Miocene Red Sea land bridges: biogeographic implications

In a large number of studies concerned with species movements between Africa and Eurasia, including the migrations of hominids out of Africa, a frequently‐cited dispersal route is across a hypothetical land bridge in the southern Red Sea, which is suggested to have emerged during glacial sea‐level lowstands. This paper, however, unequivocally demonstrates that palaeoceanographic and palaeoecological data are incompatible with the existence of Red Sea land bridges since the Miocene. The case is made by presenting the first quantitative history of water depth above the Red Sea sill for the last 470,000 years, a time period that includes the four most recent glacial–interglacial cycles, and by discussing the predictable consequences of any land bridge formation on the Red Sea sedimentary and microfossil records. The absence of post‐Miocene Red Sea land bridges has extensive implications for biogeographic models in the Afro‐Arabian region. Genetic, morphometric and palaeontological patterns reported in the literature cannot be related to dispersals over a land bridge, or in the case of marine organisms, separation of the Red Sea from the Indian Ocean by a land bridge. If such patterns in terrestrial species are only congruent with a southern Red Sea dispersal route, then they need to be considered in terms of sweepstake rafting, anthropogenic introduction, or in the particular case of the Out‐of‐Africa migration by modern humans, seafaring. The constraints imposed by our palaeoenvironmental record on biogeographic reconstructions within and around the Red Sea will hopefully encourage both the review of previous works and the preference for multidisciplinary approaches in future studies.     

Environmental changes off North Iceland during the deglaciation and the Holocene: foraminifera, diatoms and stable isotopes

A combined study of foraminifera, diatoms and stable isotopes in marine sediments off North Iceland records major changes in sea surface conditions since about 15 800 cal years (yr) BP. Results are presented from two gravity cores obtained at about 400 m water depth from two separate sedimentary basins on each side of the submarine Kolbeinsey Ridge. The chronology of the sedimentary record is based partly on AMS ¹⁴C dates, partly on the Vedde and the Saksunarvatn tephra markers, as well as the historical Hekla AD 1104 tephra. During the regional deglaciation, the planktonic foraminiferal assemblages are characterised by consistently high percentages of sinistrally coiled Neogloboquadrina pachyderma. However, major environmental variability is reflected by changes in stable isotope values and diatom assemblages. Low δ¹⁸O values indicate a strong freshwater peak as well as possible brine formation by sea-ice freezing during a pre-Bølling interval (Greenland Stadial 2), corresponding to the Heinrich 1 event. The foraminifera suggest a strong concurrent influence of relatively warm and saline Atlantic water, and both the foraminifera and the diatoms suggest mixing of cold and warm water masses. Similar but weaker environmental signals are observed during the Younger Dryas (Greenland Stadial 1) around the level of the Vedde Ash. Each freshwater peak is succeeded by an interval of severe cooling both at the beginning of the Bølling–Allerød Interstadial Complex (Greenland Interstadial 1) and during the Preboreal, presumably associated with the onset of intense deep water formatiom in the Nordic Seas. The Holocene thermal optimum, between 10 200 and about 7000 cal years (yr) BP, is interrupted by a marked cooling of the surface waters around 8200 cal yr BP. This cold event is clearly expressed by a pronounced increase in the percentages of sinistrally coiled N. pachyderma, corresponding to a temperature decrease of about 3°C. A general cooling in the area is indicated after 7000–6000 cal yr BP, both by the diatom data and by the planktonic foraminiferal data. After a severe cooling around 6000 cal yr BP, the planktonic foraminiferal assemblages suggest a warmer interval between 5500 and 4500 cal yr BP. Minor temperature fluctuations are reflected both in the foraminiferal and in the diatom data in the upper part of the record, but the time resolution of the present data is not high enough to pick up details in environmental changes through the late Holocene.     

Palaeoenvironments of deposition of Neogene laminated diatom mat deposits from the eastern equatorial Pacific from studies of benthic foraminifera (sites 844, 849, 851)

Massive sedimentation of mats of the diatom Thalassiothrix longissima forming laminated diatom mat deposits (LDM) occurred intermittently in the equatorial Pacific throughout the Neogene from at least 15 to 4.8 m.y. ago. The background deposition was otherwise calcareous nannofossil diatom ooze (NO). Benthic foraminifera have been used to reconstruct the benthic environment of deposition and the role of both surface waters (as a source of food) and bottom waters (including their corrosivity) during LDM deposition. Three LDM events were studied: Site 844 (11.4 Ma, early Tortonian), Site 849 (4.8 Ma, early Zanclian and 6.6–6.8 Ma, early Messinian). A control section of NO spanning the 4.8 Ma event was studied from Site 851. In addition, the carbonate high NO immediately preceding the 4.8 Ma event was examined in Site 849. From the foraminiferal evidence it can be shown that the LDM was deposited under conditions of normal oxygenation because infaunal taxa are present throughout and there was an increase in epifaunal taxa such as Cibicides spp. However, there was a reduction in test size probably because only those smaller than 250 μm were able to move through the restricted sediment pore spaces caused by the diatom mat meshwork. The rate of accumulation of tests is highest in LDM interval and may reflect reduced predation from macrofaunal endobenthos. Among the epifaunal taxa, Nuttallides spp. show a decrease in abundance in the LDM and even immediately preceding it. This may be linked to a greater influence of corrosive AABW. Species known to feed opportunistically on phytodetritus (Epistominella exigua and Alabaminella weddellensis) are common in both NO and LDM. The increased relative abundance of A. weddellensis in the LDM may be due to this species being smaller than E. exigua and better able to exploit the food resource within the restrictive mat meshwork. The foraminiferal results corroborate the previous interpretation that preservation of lamination in the LDM is due to the physical exclusion of macro endobenthos rather than oxygen depletion of the bottom waters.