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Invasive alien species threaten biodiversity and ecosystem structure and functioning, but incomplete assessments of their origins and temporal trends impair our ability to understand the relative importance of different factors driving invasion success. Continuous time-series are needed to assess invasion dynamics, but such data are usually difficult to obtain, especially in the case of small-sized taxa that may remain undetected for several decades. In this study, we show how micropaleontologic analysis of sedimentary cores coupled with radiometric dating can be used to date the first arrival and to reconstruct temporal trends of foraminiferal species, focusing on the alien Amphistegina lobifera and its cryptogenic congener A. lessonii in the Maltese Islands. Our results show that the two species had reached the Central Mediterranean Sea several decades earlier than reported in the literature, with considerable implications for all previous hypotheses of their spreading patterns and rates. By relating the population dynamics of the two foraminifera with trends in sea surface temperature, we document a strong relationship between sea warming and population outbreaks of both species. We conclude that the micropaleontologic approach is a reliable procedure for reconstructing the bioinvasion dynamics of taxa having mineralized remains, and can be added to the toolkit for studying invasions., peer-reviewed

The occurrence of agglutinated foraminiferal specimens belonging to the Badenian (middle Miocene) genus Colominella Popescu, 1998 was recently documented for the first time in a lower Pliocene succession of the western Mediterranean area. Direct comparisons with topotype specimens of Colominella paalzowi (Cushman 1936), sampled in the Badenian type section of Lăpugiu de Sus (Transylvania), show that the Pliocene individuals from the western Mediterranean morphologically resemble the type species C. paalzowi, but they also differ in possessing a longer biserial chamber arrangement with a higher number of internal chamber partitions, in lacking a clear early triserial stage and in having a more complex microstructure of the agglutinated wall, thereby supporting the idea that the Pliocene Mediterranean specimens represent a new, more highly evolved species. The fact that the Pliocene individuals from the Mediterranean appear to be more evolved with respect to the Badenian specimens from Paratethys represents an interesting evolutionary development of the genus Colominella that also permits the known stratigraphical and geographical range of the genus, previously limited to the middle Miocene (Badenian) of the Paratethys, to be extended.

This study updates the current distribution, range expansion and establishment status of the non-indigenous species Amphistegina lobifera Larsen, 1976 and other foraminifera that are cryptogenic in the Sicily Channel. Prior to this study, amphisteginids were reported from the Levantine Basin, the Central Mediterranean (Tunisia, Malta, Pelagian islands) and the southern Adriatic Sea. Here, we provide new records documenting a north-western expansion in the Central Mediterranean. In summer-autumn 2017 and spring-summer 2018, we collected algae and sediment samples from shallow coastal habitats along the shores of the Maltese archipelago, southern and north-western Sicily, Pantelleria and the Aegadian islands. Analysis of the foraminiferal assemblages showed that A. lobifera is effectively established around Malta and in southern/south-eastern Sicily, and has reached the oceanographic boundary between the Central and Western Mediterranean. Our results also show that the thermotolerant A. lobifera is at an advanced stage of invasion in the Sicily Channel, probably favoured by a recent rise in Mediterranean sea surface temperatures. New species distribution models are provided for the years 2040–2050 and 2090–2100, indicating that the predicted warming trend will facilitate north-westward migration of Mediterranean amphisteginids along the coast of northern Africa into the Alboran Sea, and deep into the Adriatic Sea.

Agglutinated foraminifera from surface sediments of two sites (S1 and E1) in the North Adriatic Sea were investigated in order to detect their test composition and to explore possible links with the surrounding environment. Chemical-mineralogical analyses of the agglutinated tests by scanning Electron Microscopy and Energy Dispersive Spectroscopy, suggest that the chemical composition of the test surfaces generally mirrors the one of the sea-floor sediment. Only some species, as Reophax nana and Leptohalysis scottii exhibit a clear selectivity of the agglutinated grains. In detail, specimens of R. nana from site E1, which is mainly characterized by high hydrodynamic conditions at the sea-floor, show a preferential selection of mineral grains containing high concentrations of Zircon (Zr) and Titanium (Ti) even if these elements occur in very low concentrations in the surrounding sediment. L. scottii exclusively picks mica flakes to build the test. We suggest that the compositional differences recorded in the considered agglutinated foraminiferal tests represent distinctive life strategies in order to live successfully in different environments.

This project was funded by “FAR 2012-2014” grants of the University of Pavia (Italy). The paper focuses on the occurrence of morphologically abnormal specimens of planktonic foraminifera observed over the last 550ky in IMAGES core MD 97-2114 (East of New Zealand, SW Pacific). Abnormal tests occurred throughout the entire record in all the morphospecies characterising the assemblages but were relatively rare, with percentages not exceeding 1.5% of the total assemblage. No mass abnormality events were found. A range of malformations were observed from slight deformity with smaller or overdeveloped chambers to more severe deformity, with misplaced chambers, distorted spirals or double tests forming twinned individuals. They exhibited several different categories of morphological abnormalities, even within the same sample. Test abnormalities were most abundant in the morphospecies Globorotalia inflata, Globigerina bulloides and Orbulina universa and were characterised by a long-term decreasing trend up core with an alternating % abundance pattern at the glacial to interglacial scale between MIS 14 to MIS 8, recording the highest percentages during the interglacials. Normalised total abundance and abnormal abundance curves co-varied very closely for G. bulloides and O. universa, but for G. inflata two opposing excursions were observed during MIS 13 and 6 which may be linked to water column states. Although abnormal numbers were proportionately low, there appears to be a “natural” background number of malformations in the G. inflata population through time. There was no relationship between volcanic ash production and test abnormalities. Postprint

The effects on calcite dissolution of both oceanic circulation and volcanic ash-fall were evaluated in lower bathyal sediments over the last 550 ka record from core MD 97-2114, recovered on the northern slope (depth of 1936 m, in the Pacific Deep Water, PDW) of the Chatham Rise (east of New Zealand, SW Pacific Ocean). This area has been impacted by changes in glacial/interglacial circulation and ocean chemistry as well as by the explosive volcanic activity of the Taupo Volcanic Zone. Several micro-paleontological dissolution proxies, based on planktonic foraminifera and calcareous nannofossils, were analysed in order to evaluate the calcite dissolution of the deep-sea sediments. These were compared with a couple of proxies of primary productivity (benthic foraminiferal epifaunal/infaunal ratio and δ13Cbenthic foraminifera) and the abundance of volcanic glass. The dissolution proxy data from MD 97-2114 were compared with two nearby ODP sites, ODP 1123 (3290 m deep, bathed by the lower Circumpolar Deep Water, LCDW) and ODP 1125 (1365 m deep, bathed by the Antarctic Intermediate Water, AAIW). The results suggest: (1) the calcite dissolution/preservation cycles at all three core sites show Glacial–Interglacial (G–I) periodicities that match the previously described “Pacific-style” CaCO3 cycles; (2) several short-term dissolution events do not follow this general scheme and occur following tephra deposition. The dissolution related to the tephra deposition seems to have mostly affected calcareous nannofossils, thus we hypothesise that the ash-fall induced a temporary reduction of the surface water pH (below 7.8), which affected the coccolithophores that inhabit the surface waters. (3) Other short-term dissolution events (1000 years) are unrelated to tephra deposition and are possibly driven by the slowing of deep-sea circulation and a reduced Deep Western Boundary Current (DWBC). This lead to the dominance of older, more corrosive Pacific Deep Water (PDW) flowing in to the region, resulting in coeval dissolution episodes at all three core sites (depth range from 1365 to 3290 m).

The agglutinated foraminiferal content from the last 550 kyr record of the IMAGES core MD 97-2114 (Chatham Rise, New Zealand) was analysed in order to detect the possible linkage existing between the composition of the grains forming the agglutinated tests and the deposition of tephras. The core was collected east of New Zealand, about 680 km from the active Taupo Volcanic Zone (TVZ) located on the North Island, thus it contains numerous macro- and microscopic tephra layers. Scanning Electron Microscopy and Energy Dispersive Spectroscopy analyses were carried out on entire agglutinated foraminifera as well as on sectioned specimens, sampled around and within the tephra layers. The analyses show that the studied foraminifera built structurally complex tests picking and selecting mineral and biogenic particles on the basis of their availability and abundance in the substratum, as well as their composition, size and shape. In most of the studied species, belonging to the order Textulariida, the composition of the agglutinated grains does not change when the deposition of the tephra layer strongly enriched the substratum in volcanic glass shards. Only the species Karreriella novangliae changed significantly its grain composition, mostly selecting volcanic glass fragments to cover the test surface. Nevertheless, the tephra deposition seems to influence the wall microstructure of the agglutinated tests. Textulariid specimens coming from the volcanoclastic layers have a thinner wall which is also characterised by a less abundant calcareous matrix with respect to the specimens sampled above or below the tephra layer. We hypothesise that the volcanic ash deposition probably interfered with the normal agglutinating process by causing the development of more aggressive waters at the sea floor which, in turn, could have induced carbonate dissolution. Our observations therefore suggest that the sediment type of the substratum is not the only controlling factor in the construction of the agglutinated foraminiferal test and grain selection, which appears to be species-dependent.

Over 100 cosmopolitan species of deep-sea benthic foraminifera (Extinction Group, Ext. Gp) became extinct during the late Pliocene-middle Pleistocene (3.6–0.55 Ma). Most had elongate, cylindrical tests and terminal apertures with complex modifications. This study provides new hypotheses on the functions of the morphologies that characterised the Ext. Gp and how these features could have been associated with their demise. From our functional morphological analysis we infer that: i) their elongate cylindrical or flabelliform tests, combined with fine perforations and a complex terminal apertural face are indicative of infaunal k-strategists with a low rate of metabolism; and ii) their complex apertural faces may also have been an adaptation for gathering or processing their specific phytodetrital food. We propose three alternative hypotheses for the cause of these extinctions, and where possible test them using our high resolution micropaleontological and geochemical record through the last 1.07 Ma in lower bathyal site MD 97-2114 in the SW Pacific Ocean. Hypothesis 1 is that the Ext. Gp species were unable to adapt to increased variability in the overall quantity or pulsed seasonality of the food supply to the sea floor and were out-competed by opportunistic r-strategist benthic foraminifera. This is supported by the highly variable and increasing abundance of opportunistic foraminifera at our study site during the final phase of the extinction in the mid-Pleistocene Climate Transition, MPT. We doubt however, that there was increased variability in phytoplankton productivity throughout the world's oceans sufficient to bring about the global demise of the Ext. Gp. Hypothesis 2 is that lowered pCO2 during increasingly severe MPT glacials, which coincided with the final phases of the extinction, may have caused the decline and possible loss of the Ext. Gp's phytoplankton food source. Declining pCO2 during Neogene cooling was coeval with declining relative abundance of the Ext. Gp and reticulofenestrid nannofossils, but the final demise of this latter phytoplankton group occurred slightly later than the MPT in our study site and cannot be implicated with the extinction. If this hypothesis has any validity maybe the phytoplankton group left no fossil record. Our third alternative hypothesis is that maybe our Ext. Gp had much common DNA which made them the selective target of pathogens that caused their extinction. This does not easily explain their earlier disappearance at abyssal depths than at bathyal depths in our study region, which can be accommodated by hypotheses 1 and 2.

Two sections of Epiligurian sediments (Northern Apennines, Italy) covering the chronostratigraphic interval from the Middle Eocene to the Lower Miocene were analyzed for their foraminiferal content. Several specimens belonging to 25 selected species of agglutinated foraminifera were analyzed using the energy-dispersive x-ray spectrometry (EDS) method. The collected data show how these species are selective in composition, size, and shape of the agglutinated grains and cement composition. Test microstructures can be subdivided into four groups: A relationship between test microstructure and paleobathymetric conditions for the Monte Piano Marl, Ranzano Sandstone, and Antognola Marl Formations was observed. An almost exclusive presence of agglutinated taxa showing microstructures I and II characterized the “facies varicolori” of the Monte Piano Marls (lower-middle bathyal depths, from 2000 m to 1000 m). A sharp increase in species showing microstructures III and, subordinately, IV characterized the overlying lithostratigraphic units (“facies grigia” of the Monte Piano Marls, Ranzano Sandstones and Antognola Marls) whose paleobathymetry decreases upwards (middle-upper bathyal up to slope/shelf break from 1000 to 200 m).