Heading for New Shores: Projecting Marine Distribution Ranges of Selected Larger Foraminifera

The distribution of modern symbiont-bearing larger foraminifera is confined to tropical and subtropical shallow water marine habitats and a narrow range of environmental variables (e.g. temperature). Most of today''s taxa are restricted to tropical and subtropical regions (between 30°N and 30°S) and their minimum temperature limits are governed by the 14 to 20°C isotherms. However, during times of extensive global warming (e.g., the Eocene and Miocene), larger foraminifera have been found as far north as 50°N (North America and Central Europe) as well as towards 47°S in New Zealand. During the last century, sea surface temperatures have been rising significantly. This trend is expected to continue and climate change scenarios for 2050 suggest a further increase by 1 to 3°C. We applied Species Distribution Models to assess potential distribution range changes of three taxa of larger foraminifera under current and future climate. The studied foraminifera include Archaias angulatus, Calcarina spp., and Amphistegina spp., and represent taxa with regional, superregional and global distribution patterns. Under present environmental conditions, Amphistegina spp. shows the largest potential distribution, apparently due to its temperature tolerance. Both Archaias angulatus and Calcarina spp. display potential distributions that cover currently uninhabited regions. Under climate conditions expected for the year 2050, all taxa should display latitudinal range expansions between 1 to 2.5 degrees both north- and southward. The modeled range projections suggest that some larger foraminifera may colonize biogeographic regions that so far seemed unsuitable. Archaias angulatus and Calcarina spp. also show an increase in habitat suitability within their native occurrence ranges, suggesting that their tolerance for maximum temperatures has yet not been fully exploited and that they benefit from ocean warming. Our findings suggest an increased role of larger foraminifera as carbonate producers and reef framework builders in future oceans.     

Molecular evidence for widespread occurrence of Foraminifera in soils

Environmental SSU rDNA‐based surveys are contributing to the dramatic revision of eukaryotic high‐level diversity and phylogeny as the number of sequence data increases. This ongoing revolution gives the opportunity to test for the presence of some eukaryotic taxa in environments where they have not been found using classical microscopic observations. Here, we test whether the foraminifera, a group of single‐celled eukaryotes, considered generally as typical for the marine ecosystems are present in soil. We performed foraminiferal‐specific nested PCR on 20 soil DNA samples collected in contrasted environments. Unexpectedly, we found that the majority of the samples contain foraminiferal SSU rDNA sequences. In total, we obtained 49 sequences from 17 localities. Phylogenetic analysis clusters them in four groups branching among the radiation of early foraminiferal lineages. Three of these groups also include sequences originated from previous freshwater surveys, suggesting that there were up to four independent colonization events of terrestrial and/or freshwater ecosystems by ancestral foraminifera. As shown by our data, foraminifera are a widespread and diverse component of soil microbial communities. Yet, identification of terrestrial foraminiferal species and understanding of their ecological role represent an exciting challenge for future research.     

Response of Benthic Foraminifera to Organic Matter Quantity and Quality and Bioavailable Concentrations of Metals in Aveiro Lagoon (Portugal)

This work analyses the distribution of living benthic foraminiferal assemblages of surface sediments in different intertidal areas of Ria de Aveiro (Portugal), a polihaline and anthropized coastal lagoon. The relationships among foraminiferal assemblages in association with environmental parameters (temperature, salinity, Eh and pH), grain size, the quantity and quality of organic matter (enrichment in carbohydrates, proteins and lipids), pollution caused by metals, and mineralogical data are studied in an attempt to identify indicators of adaptability to environmental stress. In particular, concentrations of selected metals in the surficial sediment are investigated to assess environmental pollution levels that are further synthetically parameterised by the Pollution Load Index (PLI). The PLI variations allowed the identification of five main polluted areas. Concentrations of metals were also analysed in three extracted phases to evaluate their possible mobility, bioavailability and toxicity in the surficial sediment. Polluted sediment in the form of both organic matter and metals can be found in the most confined zones. Whereas enrichment in organic matter and related biopolymers causes an increase in foraminifera density, pollution by metals leads to a decline in foraminiferal abundance and diversity in those zones. The first situation may be justified by the existence of opportunistic species (with high reproduction rate) that can live in low oxic conditions. The second is explained by the sensitivity of some species to pressure caused by metals. The quality of the organic matter found in these places and the option of a different food source should also explain the tolerance of several species to pollution caused by metals, despite their low reproductive rate in the most polluted areas. In this study, species that are sensitive and tolerant to organic matter and metal enrichment are identified, as is the differential sensitivity/tolerance of some species to metals enrichment.     

Spatial Patterns in the Distribution,Diversity and Abundance of Benthic Foraminifera around Moorea (Society Archipelago,French Polynesia)

Coral reefs are now subject to global threats and influences from numerous anthropogenic sources. Foraminifera, a group of unicellular shelled organisms, are excellent indicators of water quality and reef health. Thus we studied a set of samples taken in 1992 to provide a foraminiferal baseline for future studies of environmental change. Our study provides the first island-wide analysis of shallow benthic foraminifera from around Moorea (Society Archipelago). We analyzed the composition, species richness, patterns of distribution and abundance of unstained foraminiferal assemblages from bays, fringing reefs, nearshore and back- and fore-reef environments. A total of 380 taxa of foraminifera were recorded, a number that almost doubles previous species counts. Spatial patterns of foraminiferal assemblages are characterized by numerical abundances of individual taxa, cluster groups and gradients of species richness, as documented by cluster, Fisher α, ternary plot and Principal Component Analyses (PCA). The inner bay inlets are dominated by stress-tolerant, mostly thin-shelled taxa of Bolivina, Bolivinella, Nonionoides, Elongobula, and Ammonia preferring low-oxygen and/or nutrient-rich habitats influenced by coastal factors such as fresh-water runoff and overhanging mangroves. The larger symbiont-bearing foraminifera (Borelis, Amphistegina, Heterostegina, Peneroplis) generally live in the oligotrophic, well-lit back- and fore-reef environments. Amphisteginids and peneroplids were among the few taxa found in the bay environments, probably due to their preferences for phytal substrates and tolerance to moderate levels of eutrophication. The fringing reef environments along the outer bay are characterized by Borelis schlumbergeri, Heterostegina depressa, Textularia spp. and various miliolids which represent a hotspot of diversity within the complex reef-lagoon system of Moorea. The high foraminiferal Fisher α and species richness diversity in outer bay fringing reefs is consistent with the disturbance-mosaic (microhabitat heterogeneity) hypothesis.Calculations of the FORAM Index (FI), a single metric index to assess reef vitality, indicate that all fore- and most back-reef environments support active carbonate accretion and provide habitat suitability for carbonate producers dependent on algal symbiosis. Lowest suitability values were recorded within the innermost bays, an area where natural and increasing anthropogenic influences continue to impact the reefs. The presence of habitat specific assemblages and numerical abundance values of individual taxa show that benthic foraminifera are excellent recorders of environmental perturbations and good indicators useful in modern and ancient ecological and environmental studies.     

Benthic Protozoa (Foraminifera,Allogromiida) As Potential Indicator Species for the Sedimentation Record of the Azov–Black Sea Basin Bottom Deposits

Allogromiida are soft-shelled benthic foraminifera protected by thin proteinaceous shells only. They lack rigid calcareous or arenaceous structures and do not undergo fossilization. Some species of the genus Psammophaga are known to accumulate crystals of certain minerals in the cytoplasm during the life cycle. Information on benthic foraminifera (allogromiids) that inhabit the Black Sea and the Sea of Azov and accumulate mineral crystals of unknown nature inside the cells is given in the present paper for the first time. Information on these organisms is proposed to be relevant for the reconstruction of sedimentation basins that existed during the past geological epochs.     

Autogamy in Allogromia laticollaris (Foraminifera)

SYNOPSIS. The life cycles of 2 strains of Allogromia laticollaris (CSH and TPA) raised on experimentally restricted diets have been studied. Cloned cultures of the strains differed in nutritional requirements for continuous reproduction. The TPA strain was more fecund. Both strains have a basically apogamic (asexual) life cycle; only occasionally were gamonts (sexually reproducing individuals) produced. They were autogamous (self-fertilizing) and fusion of the gametes took place within the parental test.
We found in CSH clones a regular alternation of vesicularly nucleated and compactly multinucleated diploid generations. The details of the life cycle of either strain are not known well enough to be able to construct life cycle diagrams more meaningful than that of Arnold. The 2 strains studied differ in cytologic and nuclear detail from each other and from the strain studied by Arnold.     

Molecular Evidence for the Hybrid Origin of Ilex dabieshanensis (Aquifoliaceae)

Ilex, the largest genus of dioecious woody plants, is a good study system to assess the role of hybridization in speciation and evolution. Ilex dabieshanensis, a tree endemic to Dabieshan Mountains region, was initially described as a new species. Based on morphological intermediacy and sympatric distribution with its putative parental species, I. cornuta and I. latifolia, we proposed it as a natural hybrid between them. In this study, we sequenced one chloroplast intergenic spacer (trnH-psbA) and two nuclear genes (gapC and nepGS) in I. dabieshanensis and its putative parental species to test the hybrid origin hypothesis. Our results showed that there were one to two differentially fixed sequence differences between I. cornuta and I. latifolia at the two nuclear genes. Twelve of the 14 individuals of I. dabieshanensis exhibited additivity in chromatograms on these differentially fixed sites at both nuclear genes, and the remaining two exhibited additivity in chromatograms on the fixed site at only the nepGS gene. Except one haplotype of I. cornuta at the nepGS gene, all of the haplotypes of I. cornuta at the two nuclear genes were well separated from those of I. latifolia, and most haplotypes of I. dabieshanensis were shared with those of I. cornuta and I. latifolia. Phylogenetic analysis of these haplotypes was largely consistent with haplotype network analysis. I. cornuta and I. latifolia differed by two nucleotide substitutions in the chloroplast intergenic spacer, and 12 individuals of I. dabieshanensis had the same sequences as I. latifolia, while the remaining two were identical with I. cornuta. The molecular data provide convincing evidence for the hybrid origin of I. dabieshanensis and asymmetrical direction of hybridization. One haplotype of I. cornuta at the nepGS gene was nested with those of I. latifolia, indicating introgression to I. cornuta.     

Molecular data reveal high taxonomic diversity of allogromiid Foraminifera in Explorers Cove (McMurdo Sound,Antarctica)

Allogromiids are organic-walled or agglutinated, single-chambered Foraminifera, common in deep-sea and polar benthic communities. The simple forms and paucity of distinctive features make allogromiid identification difficult by traditional means. Molecular phylogenetic methods offer alternative tools for species identification and are used here to investigate allogromiid diversity. We obtained 135 partial small-subunit ribosomal DNA sequences of allogromiids collected in Explorers Cove, McMurdo Sound, Antarctica. In contrast to the 27 morphotypes identified, phylogenetic analysis revealed 49 molecular types (considered separate species) that differ by more than 5% of sequence divergence. The 49 genetic types form 28 molecular supra-groups that differ by more than 20% and probably represent distinct genera or families. Large genetic distances separating the molecular types indicate unexpectedly high taxonomic diversity. Comparison of our data with sequences of non-Antarctic allogromiids suggests that Explorers Cove species might be endemic and only distantly related to comparable northern hemisphere fauna.     

Molecular versus Taxonomic Rates of Evolution in Planktonic Foraminifera

Neogene planktonic foraminifera are among the most widely used microfossils in the study of tempo and mode of evolution. Comparisons of taxonomic rates between the two major clades in this group have shown that the nonspinose globorotaliids have undergone a significantly more rapid evolutionary turnover than the spinose globigerinids (S. M. Stanleyet al., 1988,Paleobiology14, 235–249). In order to test if similar fluctuations are observed in molecular data, we have used different methods to calculate absolute and relative rates of substitutions based on 16 partial SSU rDNA sequences from representatives of both groups. According to our data, rates of substitution are relatively constant within the globigerinids with a mean value of 4.3 subst./site/10⁹years, but vary in the globorotaliid clade with three species having a rate of about 1 subst./site/10⁹years and two species evolving much faster with rates of more than 7 subst./site/10⁹years. Assuming that the fast rates result from recent accelerations, the globorotaliids have basically much slower molecular evolutionary rates than the globigerinids, in opposition to the fossil data.     

Plastogamy in Foraminifera: Glabratella ornatissima (Cushman)

SYNOPSIS. The foraminiferan Glabratella ornatissima (Cushman) undergoes plastogamy during reproduction. In this process, 2 (or rarely more) individuals join together by their apertural sides to mutually exchange gametes. The apertural sides and internal septa are dissolved, forming a single large brood chamber. After the zygotes grow to a 2- or 3-chambered stage, the young foraminifera are liberated from the enclosing parent tests.
Scanning electron micrographs and histochemically stained thin sections indicate that the reproductively mature individuals are firmly united by an organic membrane during plastogamy. This membrane is monolamellar, less than 1 μ thick and has minute lumps on its surface. Biochemically it contains a nonsulfated acid mucopolysaccharide, which is unlike the membranes associated with CaCO₃ deposition lining the interior of foraminiferal tests. The young foraminifera liberate themselves by secretion of an enzyme which degrades the membrane. The mechanism of plastogamy is interpreted to be an adaptation to life in turbulent waters that would decrease the chance of gamete union or zygote survival were the gametes released and fertilized freely.     

Ontogeny and phylogeny of Upper Cenomanian rotaliporids (Foraminifera)

The fossil record of planktonic foraminifera is a key source of data on the evolution of marine plankton. One of the most distinctive groups of Cretaceous foraminifera, the rotaliporids, widely used as a stratigraphic index, has always been considered to be a monophyletic clade. New data on the coiling direction and persistent morphological features of the late rotaliporids from the Upper Cenomanian of the Western Interior Seaway, USA, and the Vocontian Basin of southeast France is used as a phylogenetic proxy. Dealing with key morphological features, the coiling pattern of these keeled morphotypes proves that the rotaliporids group is polyphyletic and composed of Thalmanninella, that displays a dextral-coiling preference, and Rotalipora s.s., that have a proportionate-coiling mode. The stratigraphically youngest rotaliporids with keels co-occur with globular forms; and all morphologies transitional between these morphotypes are observed. The ontogenetic relationships between them are investigated, indicating that loss of the keel was a selective advantage that enabled those rotaliporids to remain in the surface water, thereby avoiding the expansion of the oxygen minimum zone. Two species are observed: Thalmanninella multiloculata and Rotalipora planoconvexa. These species are interpreted as having arisen by neoteny from Thalmanninella greenhornensis and Rotalipora cushmani respectively.     

Quantitative Studies on the Growth of Allogromia laticollaris (Foraminifera)

SYNOPSIS. The growth and reproduction of Allogromia laticollaris was studied. More schizozoites were generally produced in mixtures of food organisms than on single algal foods. In the presence of moderate numbers of bacteria, cultures with Phaeodactylum tricornutum, Chlorococcum sp., Nannochloris sp., and an unidentified chlorophyte (BL-1), added singly, were also highly productive. Schizogony was the dominant asexual form of reproduction. Binary fission and cytotomy also occurred in bacterized otherwise unfed controls. ³⁵S and ³²P are convenient labels for measuring growth of A. laticollaris when introduced into the system in the range of 1 × 10⁴ - 1 × 10⁵ dpm/ml (³²P specific activity ～ 2.03 MCi/g; ³⁵S specific activity ～ 95 μCi/g). Small allogromiids grew faster than did larger ones. By means of the Taylor series modification of the classical least-squares method, a continuous life-cycle representation was calculated for A. laticollaris for the conditions of the experiment. Four points of cell volume growth were maxima for reproduction: 1.0 × 10⁷μ per organism for curve I; 2.2 × 10⁷μ³ and 1.2 × 10⁷μ³ for curve II; and 6.7 × 10⁷μ³ for curve III.     

Molecular Evidence for Parallel Evolution of Adaptive Syndromes in Fig-BreedingLissocephala(Drosophilidae)

AfrotropicalLissocephala(Drosophilidae) breed strictly in syconia (figs) ofFicus(Moraceae) and have accordingly evolved specific features including modified female and eggshell morphologies, ovipositing, larval foraging, and mating behaviors. These various traits may exist as two or three alternative states. Each species displays a specific suite of traits so closely coordinated with one another that alternative states of the overall suites of traits can be seen as “adaptive syndromes.” Three clear-cut adaptive syndromes can be recognized while two taxonomic lineages (junctaandsanuspecies groups) are traditionally accepted on the basis of male terminalia. A crucial evolutionary question results from the consideration that ecological clusters and taxonomic groups have conflicting compositions: the three syndromes are found in thejunctagroup while two of them occur in thesanugroup. To resolve this conflict, we present molecular data which provide a robust phylogeny: mitochondrial DNA (12S + 16S ribosomal DNA and cytochromeb) sequence data are in agreement with one another regardless of the algorithm used. All molecular data consistently support male terminalia dichotomy. Such a level of consistency unambiguously indicates that parallel evolution of adaptive syndromes occurred. Thus, homoplasy may affect morphological and behavioral traits concomitantly when these are involved in a network of functional relationships.     

Molecular Evidence for Multiple Origins of the European Spined Loaches (Teleostei,Cobitidae)

We present a phylogenetic investigation of the Northern Clade, the major monophyletic clade within the freshwater fish family Cobitidae, one of the most prominent families of freshwater fishes found in Asian and European waters. Phylogenetic reconstructions based on the cytochrome b and RAG-1 genes show the genera Microcobitis, Sabanejewia, Koreocobitis and Kichulchoia as monophyletic groups. These reconstructions also show a Cobitis sensu lato and a Misgurnus sensu lato group. The Cobitis sensu lato group includes all species of Cobitis, Iksookimia, Niwaella and Kichulchoia, while the Misgurnus sensu lato group includes Misgurnus, Paramisgurnus and Koreocobitis. Although the monophyly of both the Cobitis sensu lato and Misgurnus sensu lato groups is supported, relationships within the groups are incongruent with current generic definitions. The absence of monophyly of most genera included in the Cobitis sensu lato group (Cobitis, Iksookimia and Niwaella) or their low genetic differentiation (Kichuchoia) supports their consideration as synonyms of Cobitis. Molecular phylogenies indicate that the Asian species of Misgurnus experienced a mitochondrial introgression from a lineage of Cobitis. We also find two nuclear haplotypes in the same Cobitis species from the Adriatic area that, in the absence of morphological differentiation, may indicate molecular introgression. Most lineages within the Northern Clade consist of species found in East Asia. However, some lineages also contain species from Europe and Asia Minor. The phylogenetic relationships presented here are consistent with previous studies suggesting an East Asian origin of the Northern Clade. According to the current distributions and phylogenetic relationships of the Misgurnus sensu lato and Cobitis clade lineages, particularly of M. fossilis and C. melanoleuca, the range expansion of East Asian species into Europe was most likely via Siberia into Northern and Central Europe. Phylogenetic analyses also show that the Cobitis sensu lato group consists of two clear subgroups (I and II), each presenting geographical differences. Subgroup I is distributed exclusively in East Asian drainages with an Eastern European offshoot (C. melanoleuca), whereas Subgroup II includes species widespread throughout Europe (including the Mediterranean), Asia Minor, the Black Sea and the Caucasus, with some lineages related to species restricted to East Asia.     

Molecular Evidence for a Deep Clade of Dunnarts (Marsupialia: Dasyuridae: Sminthopsis)

Sminthopsis is the most speciose genus of living dasyurid marsupials and, along with its close relatives Antechinomys and Ningaui, constitutes the clade Sminthopsini. Phylogenetic relationships among the 23 species in this clade have been the subject of much morphological and molecular investigation, including a recent integration of penis morphology (in Sminthopsis) with molecular systematics. Several phylogenetic issues remain open, however, including the monophyly of Sminthopsis and branching order among early sminthopsin lineages. In this study, we revisit sminthopsin systematics with an expanded molecular data set, including new DNA sequences from mitochondrial (valine transfer-RNA and 16S ribosomal RNA) and nuclear (interphotoreceptor retinoid binding protein and beta-fibrinogen) loci, along with previously published sequences of cytochrome b, 12S ribosomal RNA, control region, and protamine P1. Our results again fail to establish the monophyly of Sminthopsis, but do provide a clearer resolution of early sminthopsin branching. Specifically, our phylogeny suggests three major groups of Sminthopsis species: S. longicaudata (perhaps the sister of Antechinomys); the Macroura species group of previous authors (S. crassicaudata, S. macroura, S. virginiae, S. douglasi, and S. bindi); and the remaining 13 species allied with the Murina species group. Our results depart from previous molecular findings by reuniting S. ooldea with the Murina group, while resolving S. psammophila as sister to the hairy-footed dunnarts (S. hirtipes and S. youngsoni). We suggest that this conflict traces to anomalous phylogenetic signal in previously published cytochrome b sequences. Penis morphology maps reasonably well onto our phylogeny, requiring parallel origination of only one of the ten morphotypes described for Sminthopsis.