TEST FUSION IN ADULT FORAMINIFERA: A REVIEW WITH NEW OBSERVATIONS OF AN EARLY EOCENE SPECIMEN
In foraminifera, so-called "double tests" usually arise due to abnormal growth originating mainly from twinning, but may also be caused by irregularities in the early chambers and by regeneration after test injury that modifies the direction of growth. A fourth cause of double tests has only rarely been reported: the fusion of the tests of two adult individuals. We studied an early Eocene double test consisting of two adult individuals that fused after an extended period of independent growth. The specimen was studied using computed tomography with micrometric resolution (micro-CT) that allowed bi- and three-dimensional visualization of the internal structure. Before fusion each individual test had 30-36 chambers, which, by comparison with growth rates in recent nummulitids, implies at least three months of independent growth. After fusion, the compound test grew in two spirals that fused after about one whorl and then continued in a single spiral. To fuse their tests, either adult individuals have to be forced to do so or the allorecognition (ability to distinguish between self and another individual) mechanisms must fail. A possible explanation for the merged tests in this study is forced fusion in attached individuals after surviving ingestion and digestion by a metazoan. Alternatively, environmental stress could lead to a failure of allorecognition mechanisms and/or foraminiferal motility. Once fused, subsequent growth seems to be determined mainly by the relative orientation of individual tests. In any case, the frequency in which adult fusion occurs remains unknown.
PALEOBIOLOGICAL APPLICATIONS OF THREE-DIMENSIONAL BIOMETRY ON LARGER BENTHIC FORAMINIFERA: A NEW ROUTE OF DISCOVERIES
Four specimens of larger benthic foraminifera (the Recent and , and the phylogenetically related Paleogene and ) were investigated by X-ray tomography. The resulting three-dimensional measurements enabled a comprehensive, quantitative study of shell morphology to interpret cell growth without specific shell preparation and/or destruction. After segmentation and extraction of all scanned lumina, the following characters were measured on all chambers of each specimen: chamber volume, septal distance, chamber height, and chamber width. The sequence of chamber lumina follows either a logistic function (, ), where the deceleration in growth rate of the latest chambers could mark the onset of reproduction, or it can be modeled by a series of stepwise functions with differing constants (). Variations around the growth model are either periodic, following external cycles, or random as expressed by abrupt deviations. Therefore, they may reflect the response of the cell to environmental changes in terms of cyclic changes (e.g., seasonality) or single events (e.g., predator attack). Correlations between chamber volume and the other chamber parameters show that septal distance always matches the sequence in chamber volume and can therefore be used as a proxy for environmental analyses in both growth models. Chamber height and width often remain constant around their function and rarely deviate drastically to accommodate the needed lumen for retaining test size and shape. Chamber width may vary according to chamber volume in involute specimens, whereas both chamber height and width correlate with volume in those tests following an Archimedean spiral. X-ray-tomography shows particular promise in determining which parameters that can be assessed routinely in two dimensions primarily reflect environmental conditions vs. parameters best used for taxonomic identification and for systematic lineage reconstruction.
AXIALLY ORIENTED SECTIONS OF NUMMULITIDS: A TOOL TO INTERPRET LARGER BENTHIC FORAMINIFERAL DEPOSITS
The "critical shear velocity" and "settling velocity" of foraminiferal shells are important parameters for determining hydrodynamic conditions during deposition of banks. These can be estimated by determining the size, shape, and density of nummulitid shells examined in axial sections cut perpendicular to the bedding plane. Shell size and shape can be determined directly from the shell diameter and thickness, but density must be calculated indirectly from the thin section. Calculations using the half-tori method approximate shell densities by equalizing the chamber volume of each half whorl, based on the half whorl's lumen area and its center of gravity. Results from this method yield the same lumen volumes produced empirically by micro-computed tomography. The derived hydrodynamic parameters help estimate the minimum flow velocities needed to entrain nummulitid tests and provide a potential tool to account for the nature of their accumulations.