Phytogeography of New Guinean orchids: richness, turnover, collection density

The aims of this study were (1) to assess the spatial distribution of orchid species richness in New Guinea, and (2) to examine patterns of species turnover in the orchid community through phytogeographical regionalization. We aimed to achieve these goals using botanical collection records, species distribution models (SDMs) and partitioning around medoids (PAM) clustering.

Middle Eocene to early Oligocene planktonic and benthic foraminifers from ODP Hole 125-786A

Drilling at Site 786, located in the center of the Izu-Bonin forearc basin, penetrated an apparently continuous section of middle Eocene/lower Oligocene volcaniclastic breccias and nannofossil oozes. Planktonic foraminiferal faunas underwent a gradual transition from relatively high-diversity middle Eocene through late Eocene tropical or warm-water assemblages to a cooler-water, less diverse assemblage during the early Oligocene. In the cosmopolitan benthic foraminiferal faunas, the major transition occurred during the early late Eocene. Middle Eocene benthic assemblages resembling the bathyal 'Lenticulina' fauna (characterized by Osangularia mexicana, Cibicidoides eocaenus, and several buliminid species) changed to an upper Eocene abyssal 'Globocassidulina subglobosa' fauna (characterized by Cibicidoides praemundulus, Globocassidulina subglobosa, Gyroidinoides girardanus, Oridorsalis umbonatus, and Siphonodosaria aculeata). Even though no large, abrupt faunal changes appear to have been associated with the assumed Eocene/Oligocene boundary, benthic species turnover continued through the late Eocene and into the early Oligocene. This resulted in a slightly lower diversity early Oligocene fauna dominated by three species: Laevidentalina sp., Bulimina jarvisi, and Gyroidinoides girardanus. The progression from a middle Eocene bathyal 'Lenticulina' fauna, rather than an abyssal 'Nuttallides truempyi' fauna, to an abyssal 'Globocassidulina subglobosa' fauna during the early late Eocene, suggests that a bathymetric deepening occurred at Site 786. Increased water depths may have resulted from tectonic subsidence.

Strontium isotopic ratios in selected microfossils from ODP Holes 119-738C and 120-747A

The Maastrichtian and Danian intervals of Ocean Drilling Program (ODP) Hole 738C contain numerous microfossils above the level of their putative extinction, suggesting either (1) persistence of local communities long after species turnover occurred across the rest of the globe or (2) large-scale reworking. These interpretations have very different paleoenvironmental implications, but discriminating between them has proved difficult. To test the competing hypotheses, we measured the 87Sr/86Sr ratios of taxon-specific separates from a number of samples and compared these values both to each other and to expected seawater values at the time of deposition. Our results indicate extensive and pervasive reworking throughout Maastrichtian and lower Danian strata in ODP Hole 738C. We estimate that up to 30% of the mass of foraminifers in any sample can be contributed by individuals that have been reworked.

Benthic foraminifera extinctions in the Cenozoic record

In the late Pliocene-middle Pleistocene a group of 95 species of elongate, cylindrical, deep-sea (lower bathyal-abyssal) benthic foraminifera became extinct. This Extinction Group (Ext. Gp), belonging to three families (all the Stilostomellidae and Pleurostomellidae, some of the Nodosariidae), was a major component (20-70%) of deep-sea foraminiferal assemblages in the middle Cenozoic and subsequently declined in abundance and species richness before finally disappearing almost completely during the mid-Pleistocene Climatic Transition (MPT). So what caused these declines and extinction? In this study 127 Ext. Gp species are identified from eight Cenozoic bathyal and abyssal sequences in the North Atlantic and equatorial Pacific Oceans. Most species are long-ranging with 80% originating in the Eocene or earlier. The greatest abundance and diversity of the Ext. Gp was in the warm oceanic conditions of the middle Eocene-early Oligocene. The group was subjected to significant changes in the composition of the faunal dominants and slightly enhanced species turnover during and soon after the rapid Eocene-Oligocene cooling event. Declines in the relative abundance and flux of the Ext. Gp, together with enhanced species loss, occurred during middle-late Miocene cooling, particularly at abyssal sites. The overall number of Ext. Gp species present began declining earlier at mid abyssal depths (in middle Miocene) than at upper abyssal (in late Pliocene-early Pleistocene) and then lower bathyal depths (in MPT). By far the most significant Ext. Gp declines in abundance and species loss occurred during the more severe glacial stages of the late Pliocene-middle Pleistocene. Clearly, the decline and extinction of this group of deep-sea foraminifera was related to the function of their specialized apertures and the stepwise cooling of global climate and deep water. We infer that the apertural modifications may be related to the method of food collection or processing, and that the extinctions may have resulted from the decline or loss of their specific phytoplankton or prokaryote food source, that was more directly impacted than the foraminifera by the cooling temperatures.

Dinoflagellate cyst abundances at DSDP Site 94-607 from the North Atlantic (Appendix B)

The climatic deterioration related to the onset of Northern Hemisphere glaciations (circa 2.52 Ma BP) must have lead to reorganization and relocation of species associations and may have enhanced species turnover. The present study investigates how this deterioration affects the dinoflagellate cyst and acritarch assemblages from two locations, DSDP Site 607 (North Atlantic) and the Singa section (southern Italy). The records from these locations cover the interval from 2.8 to 2.2 Ma with at least a 5 ka resolution and they have been correlated to the Milankovitch periodicities on a cycle to cycle basis by means of integrated high resolution stable isotope, calcium carbonate, foraminiferal, palynological and magnetostratigraphical datasets. In the present study this high resolution stratigraphic framework is used for a detailed correlation of events occurring in each of the depositional sequences. It also enables further assessment of the palaeoenvironmental preferences of some dinoflagellate cyst forms. Comparison of the two palynological records reveals a close correspondence in the timing of major assemblage changes and extinction events, confirming their Milankovitch cycle based correlation. A close link between periods of Northern Hemisphere cooling (at oxygen isotope stages 110, 104 and 100-96) and increased dinoflagellate cyst turnover appears to be present for both DSDP Site 607 and the Singa section. The turnover events can also be recognized in the records of planktic foraminifera and calcamous nannoplankton. Comparison of the Singa section with Site 607 and with other time equivalent marine palynological data sets, shows that some oceanic taxa respond similarly over a large area. The biostratigraphical implications are discussed. Notably the last occurrence of Invertocystu lucrymosa appears to be a valuable marker for isotope stage 110 in the Mediterranean and North Atlantic.

Multi-temporal mapping of seagrass cover, species and biomass of the Eastern Banks, Moreton Bay, Australia, with links to shapefiles

The spatial and temporal dynamics of seagrasses have been studied from the leaf to patch (100 m**2) scales. However, landscape scale (> 100 km**2) seagrass population dynamics are unresolved in seagrass ecology. Previous remote sensing approaches have lacked the temporal or spatial resolution, or ecologically appropriate mapping, to fully address this issue. This paper presents a robust, semi-automated object-based image analysis approach for mapping dominant seagrass species, percentage cover and above ground biomass using a time series of field data and coincident high spatial resolution satellite imagery. The study area was a 142 km**2 shallow, clear water seagrass habitat (the Eastern Banks, Moreton Bay, Australia). Nine data sets acquired between 2004 and 2013 were used to create seagrass species and percentage cover maps through the integration of seagrass photo transect field data, and atmospherically and geometrically corrected high spatial resolution satellite image data (WorldView-2, IKONOS and Quickbird-2) using an object based image analysis approach. Biomass maps were derived using empirical models trained with in-situ above ground biomass data per seagrass species. Maps and summary plots identified inter- and intra-annual variation of seagrass species composition, percentage cover level and above ground biomass. The methods provide a rigorous approach for field and image data collection and pre-processing, a semi-automated approach to extract seagrass species and cover maps and assess accuracy, and the subsequent empirical modelling of seagrass biomass. The resultant maps provide a fundamental data set for understanding landscape scale seagrass dynamics in a shallow water environment. Our findings provide proof of concept for the use of time-series analysis of remotely sensed seagrass products for use in seagrass ecology and management.

Neogene planktonic foraminifera of ODP Site 130-806

The Neogene biostratigraphy presented here is based on the study of 230 samples through 737 m of pelagic sediment in Hole 806B. Sediment accumulation is interrupted only once in the uppermost lower Miocene (Zone N6), apparently coincident with a widespread deep-sea hiatus. Preservation of planktonic foraminifers through the section ranges from good to moderately poor. One hundred and ten species of planktonic foraminifers were identified; taxonomic notes on most species are included. All of the standard low-latitude Neogene foraminiferal zones are delineated, with the exceptions of Zones N8 and N9 because of a high first occurrence of Orbulina, and Zones N18 and N19 because of a high first occurrence of Sphaeroidinella dehiscens. Good agreement exists between the published account of the variation in planktonic foraminiferal species richness and the rates of diversification and turnover, and measurements of these evolutionary indexes in the record of Hole 806B. The global pattern of change in tropical/transitional species richness is paralleled in Hole 806B, with departures caused by either ecological conditions peculiar to the western equatorial Pacific or by inexactness in the estimation of million-year intervals in Hole 806B. Temporal changes in the relative abundance of taxa in the sediment assemblages, considered in light of their depth habitats, reveal a detailed picture of historical change in the structure of the upper water column over the Ontong Java Plateau. The dominance of surface dwellers (Paragloborotalia kugleri, P. mayeri, Dentoglobigerina altispira, Globigerinita glutinata, and Globigerinoides spp.) throughout the lower and middle Miocene is replaced by a more equitable distribution of surface (D. altispira and Globigerinoides spp.), intermediate (Globorotalia menardii plexus), and deep (Streptochilus spp.) dwellers in the late Miocene, following the closing of the Indo-Pacific Seaway and the initiation of large-scale glaciation in the Antarctic. The shoaling of the thermocline along the equator engendered by these climatic and tectonic events persisted through the Pliocene, when initial increases in the abundance of a new set of shallow, intermediate, and deep dwelling species of planktonic foraminifers coincide with the closing of the Panamanian Seaway.

Calcareous nannofossil species richness across the Paleocene-Eocene Thermal Maximum

The Paleocene-Eocene Thermal Maximum (PETM, ~5 million years ago) was an interval of global warming and ocean acidification attributed to rapid release and oxidation of buried carbon. We show that the onset of the PETM coincided with a prominent increase in the origination and extinction of calcareous phytoplankton. Yet major perturbation of the surface-water saturation state across the PETM was not detrimental to the survival of most calcareous nannoplankton taxa and did not impart a calcification or ecological bias to the pattern of evolutionary turnover. Instead, the rate of environmental change appears to have driven turnover, preferentially affecting rare taxa living close to their viable limits.

Radiolarian faunal turnover across the Oligocene/Miocene boundary in the equatorial Pacific Ocean

The global warming trend of the latest Oligocene was interrupted by several cooling events associated with Antarctic glaciations. These cooling events affected surface water productivity and plankton assemblages. Well-preserved radiolarians were obtained from upper Oligocene to lower Miocene sediments at Ocean Drilling Program (ODP) Leg 199 Sites 1218 and 1219 in the equatorial Pacific, and 110 radiolarian species were identified. Four episodes of significant radiolarian faunal changes were identified: middle late Oligocene (27.5 to 27.3 Ma), latest Oligocene (24.4 Ma), earliest Miocene (23.3 Ma), and middle early Miocene (21.6 Ma). These four episodes approximately coincide with increases and decreases of biogenic silica accumulation rates and increases in delta18O values coded as "Oi" and "Mi" events. These data indicate that Antarctic glaciations were associated with change of siliceous sedimentation patterns and faunal changes in the equatorial Pacific. Radiolarian fauna was divided into three assemblages based on variations in radiolarian productivity, species richness and the composition of dominant species: a late Oligocene assemblage (27.6 to 24.4 Ma), a transitional assemblage (24.4 to 23.3 Ma) and an early Miocene assemblage (23.3 to 21.2 Ma). The late Oligocene assemblage is characterized by relatively high productivity, low species richness and four dominant species of Tholospyris anthophora, Stichocorys subligata, Lophocyrtis nomas and Lithelius spp. The transitional assemblage represents relatively low values of productivity and species richness, and consists of three dominant species of T. anthophora, S. subligata and L. nomas. The characteristics of the early Miocene assemblage are relatively low productivity, but high species richness. The two dominant species present in this assemblage are T. anthophora and Cyrtocapsella tetrapera. The most significant faunal turnover of radiolarians is marked at the boundary between the transitional/early Miocene assemblages.

Lipid, carbon and nitrogen content and fatty acid composition of Temora longicornis females and different food species during the experiment

Temora longicornis, a dominant calanoid copepod species in the North Sea, is characterised by low lipid reserves and high biomass turnover rates. To survive and reproduce successfully, this species needs continuous food supply and thus requires a highly flexible digestive system to exploit various food sources. Information on the capacity of digestive enzymes is scarce and therefore the aim of our study was to investigate the enzymatic capability to respond to quickly changing nutritional conditions. We conducted two feeding experiments with female T. longicornis from the southern North Sea off Helgoland. In the first experiment in 2005, we tested how digestive enzyme activities and enzyme patterns as revealed by substrate SDS-PAGE (sodium dodecylsulfate polyacrylamide gel electrophoresis) responded to changes in food composition. Females were incubated for three days fed ad libitum with either the heterotrophic dinoflagellate Oxyrrhis marina or the diatom Thalassiosira weissflogii. At the beginning and at the end of the experiment, copepods were deep-frozen for analyses. The lipolytic enzyme activity did not change over the course of the experiment but the enzyme patterns did, indicating a distinct diet-induced response. In a second experiment in 2008, we therefore focused on the enzyme patterns, testing how fast changes occur and whether feeding on the same algal species leads to similar patterns. In this experiment, we kept the females for 4 days at surplus food while changing the algal food species daily. At day 1, copepods were offered O. marina. On day 2, females received the cryptophycean Rhodomonas baltica followed by T. weissflogii on day 3. On day 4 copepods were again fed with O. marina. Each day, copepods were frozen for analysis by means of substrate SDS-PAGE. This showed that within 24 h new digestive enzymes appeared on the electrophoresis gels while others disappeared with the introduction of a new food species, and that the patterns were similar on day 1 and 4, when females were fed with O. marina. In addition, we monitored the fatty acid compositions of the copepods, and this indicated that specific algal fatty acids were quickly incorporated. With such short time lags between substrate availability and enzyme response, T. longicornis can successfully exploit short-term food sources and is thus well adapted to changes in food availability, as they often occur in its natural environment due seasonal variations in phyto- and microzooplankton distribution.

Planktic foraminiferal assemblage turnover at the Cretaceous-Tertiary boundary

Three uppermost Cretaceous through basal Paleocene stratigraphic sequences are examined for planktic foraminiferal assemblage stability and temporal succession patterns. These sequences are at mid-latitude South Atlantic DSDP Site 528, then-equatorial Pacific DSDP Site 577 and the Tethyan shelf Ben Gurion section of the Negev, Israel. In order to better estimate biogeographic patterns and habitat preferences, the results of these analyses are compared to previous Cretaceous biogeographic studies and to previous analyses of Cretaceous-Tertiary (K/T) boundary shelf and epicontinental sections. Results indicate that immediately following the K/T boundary, the examined epicontinental and open-ocean sites were exploited primarily by previously epicontinental planktic foraminiferal assemblages. This pattern of K/T boundary assemblage dominance suggests the geologically instantaneous break-down of Late Cretaceous epicontinental and open-ocean biogeographic provincialization. This shift in open-ocean foraminiferal assemblages is not consistent with models of nonselective K/T boundary extinctions, but is consistent with models of extinction resistence and offshore expansion of nearshore taxa. The re-establishment of stable biogeographic differences between open-ocean and epicontinental planktic foraminiferal assemblages occurs by the basal Parvularugoglobigerina eugubina Zone. At open-ocean sites 528 and 577 and the outershelf Ben Gurion section, P0 and P. eugubina Zone faunal records are marked by a pronounced alternation between Paleocene biserial- and non-biserial-dominated assemblages, This alternation appears strongly damped at shelf and epicontinental sections previously examined. The first appearance and peak magnitude of abundant earliest Paleocene trochospiral forms (Parvularugoglobigerina, Eoglobigerina, Morozovella, Globoconusa) also vary from site to site and may depend closely on levels of primary carbonate productivity.

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.

Planktic foraminiferal turnover across the Paleocene-Eocene transition in the Bay of Biscay, North Atlantic

Planktic foraminifera across the Paleocene-Eocene transition at DSDP Site 401 indicate that the benthic foraminiferal mass extinction occurred within Subzone P 6a of Berggren and Miller (1988), or PS of Berggren et al. (1995) and coincident with a sudden 2.0? excursion in 6r3C values. The benthic foraminiferal extinction event (BFEE) and Sr3C excursion was accompanied by a planktic foraminiferal turnover marked by an influx of warm water species (Morozovella and Acarinina), a decrease in cooler water species (Subbotina), a sudden short-term increase in low oxygen tolerant taxa (Chiloguembelina), and no significant species extinctions. These faunal changes suggest climatic warming, expansion of the oxygen minimum zone, and a well stratified ocean water column. Oxygen isotope data of the surface dweller M. subbotina suggest climate warming beginning with a gradual 0.5? decrease in delta180 in the 175 cm preceding the benthic foraminiferal extinction event followed by a sudden decrease of 1? (4°C) at the BFEE. The delta13C excursion occurred over 27 cm of sediment and, assuming constant sediment accumulation rates, represents a maximum of 23 ka. Recovery to pre-excursion delta13C values occurs within 172 cm, or about 144 ka. Climate cooling begins in Subzone P 6c as indicated by an increase in cooler water subbotinids and acarininids with rounded chambers and a decrease in warm water morozovellids.

Fecal pellet and egg production rates of copepod species at varying suspended sediment concentrations

We investigated the effect of suspended sediments on the vital rates of the copepods Calanus finmarchicus, Pseudocalanus sp. and Metridia longa in a Greenland sub-Arctic fjord. The fjord had a gradient of suspended particulate matter (SPM) with high concentrations (>50 mg/L) in the inner fjord due to glacial melt water runoff. Laboratory experiments showed that when feeding on the diatom Thalassiosira weissflogii specific ingestion rates were low at high concentrations of suspended sediment for C. finmarchicus (>20 mg/L) and Pseudocalanus sp. (>50 mg/L), while no effect was found for M. longa. For C. finmarchicus, a relatively constant fecal pellet production (FPP) and fecal pellet volume suggested ingestion of sediment, which probably led to reduction in egg production rates (EPRs) at high sediment concentrations. For Pseudocalanus sp., FPP decreased with increasing sediment concentrations, while no effect was observed on EPR. No significant difference was observed in FPP for M. longa feeding on the diatom T. weissflogii compared to the ciliate Strombidium sulcatum. The study shows that high sediment concentrations influence the capability of carbon turnover in C. finmarchicus and Pseudocalanus sp., while M. longa appears to be more tolerant to high sediment loads. Therefore, high concentrations of SPM could potentially influence the species composition of glacially influenced fjords.