induction of programmed cell death provides a more direct link to study a Red Queen-like coevolutionary arms race at least between the coccolithoviruses and diploid organism. Evolution and diversity Coccolithophores are members of the clade Haptophyta, which is a sister clade to Centrohelida, which are both in Haptista. The oldest known coccolithophores are known from the Late Triassic, around the Norian-Rhaetian boundary. Diversity steadily increased over the course of the Mesozoic, reaching its apex during the Late Cretaceous. However, there was a sharp drop during the Cretaceous-Paleogene extinction event, when more than 90% of coccolithophore species became extinct. Coccoliths reached another, lower apex of diversity during the Paleocene-Eocene thermal maximum, but have subsequently declined since the Oligocene due to decreasing global temperatures, with species that produced large and heavily calcified coccoliths most heavily affected. Coccolithophore shells Exoskeleton: coccospheres and coccoliths Each coccolithophore encloses itself in a protective shell of coccoliths, calcified scales which make up its exoskeleton or coccosphere. The coccoliths are created inside the coccolithophore cell and while some species maintain a single layer throughout life only producing new coccoliths as the cell grows, others continually produce and shed coccoliths. Composition The primary constituent of coccoliths is calcium carbonate, or chalk. Calcium carbonate is transparent, so the organisms' photosynthetic activity is not compromised by encapsulation in a coccosphere. Formation Coccoliths are produced by a biomineralization process known as coccolithogenesis. Generally, calcification of coccoliths occurs in the presence of light, and these scales are produced much more during the exponential phase of growth than the stationary phase. Although not yet entirely understood, the biomineralization process is tightly regulated by calcium signaling. Calcite formation begins in the golgi complex where protein templates nucleate the formation of CaCO3 crystals and complex acidic polysaccharides control the shape and growth of these crystals. As each scale is produced, it is exported in a Golgi-derived vesicle and added to the inner surface of the coccosphere. This means that the most recently produced coccoliths may lie beneath older coccoliths. Depending upon the phytoplankton's stage in the life cycle, two different types of coccoliths may be formed. Holococcoliths are produced only in the haploid phase, lack radial symmetry, and are composed of anywhere from hundreds to thousands of similar minute (ca 0.1 μm) rhombic calcite crystals. These crystals are thought to form at least partially outside the cell. Heterococcoliths occur only in the diploid phase, have radial