Elasmobranchs are characterised by the presence of placoid scales on their skin. These scales, structurally homologous to gnathostome teeth, are thought to have various ecological functions related to drag reduction, predator defense or abrasion reduction. Some scales, particularly those present in the ventral area, are also thought to be functionally involved in the transmission of bioluminescent light in deep-sea environments. In the deep parts of the oceans, elasmobranchs are mainly represented by squaliform sharks. This study compares ventral placoid scale morphology and elemental composition of more than thirty deep-sea squaliform species. Scanning Electron Microscopy and Energy Dispersive X-ray spectrometry, associated with morphometric and elemental composition measurements were used to characterise differences among species. A maximum likelihood molecular phylogeny was computed for 43 shark species incuding all known families of Squaliformes. Character mapping was based on this phylogeny to estimate ancestral character states among the squaliform lineages. Our results highlight a conserved and stereotypical elemental composition of the external layer among the examined species. Phosphorus-calcium proportion ratios (Ca/P) slightly vary from 1.8-1.9, and fluorine is typically found in the placoid scale. By contrast, there is striking variation in shape in ventral placoid scales among the investigated families. Character-mapping reconstructions indicated that the shield-shaped placoid scale morphotype is likely to be ancestral among squaliform taxa. The skin surface occupied by scales appears to be reduced in luminous clades which reflects a relationship between scale coverage and the ability to emit light. In luminous species, the placoid scale morphotypes are restricted to pavement, bristle-and spine-shaped except for the only luminescent somniosid, Zameus squamulosus, and the dalatiid Mollisquama mississippiensis. These results, deriving from an unprecedented sampling, show extensive morphological diversity in placoid scale shape but little variation in elemental composition among Squaliformes.

Bioluminescence is a common ecological trait among many marine organisms, including three shark families: Etmopteridae, Dalatiidae, and Somniosidae. The kitefin shark, Dalatias licha (Bonnaterre, 1788), from the Dalatiidae family is the largest known luminous vertebrate. This study compares the light organ ultrastructure of D. licha with that of Etmopterus spinax, the type species of Etmopteridae, to gain a deeper understanding of the light emission process and its evolutionary conservation within shark families. The ultrastructure of D. licha’s photophores and the morphological changes that occur after hormonal stimulation (via melatonin and α-MSH, which stimulate or inhibit the bioluminescence, respectively) were examined. The photophores consist of a spherical pigmented sheath surrounding a unique, regionalized light-emitting cell (photocyte). The photocyte’s basal area contains a specific area filled with granular inclusions that resemble the glowon-type microsources of E. spinax, suggesting that this area is the intracellular site of light emission. An acidophilic secretion, not present in Etmopteridae, is also observed within the granular area and may be involved in photogenesis. The ultrastructure analysis reveals no lens cells or reticular layer, unlike in Etmopteridae photophores, indicating a simpler organization in Dalatiidae photophores. Melatonin stimulation causes the removal of pigments from the photophore-associated melanophores and an increase in the granular inclusion diameter and coverage in the granular area, further showing that this last area is the potential site of light emission, while α-MSH stimulation causes the extension of the melanophore pigments and a decrease in the granular inclusion diameter and coverage. These results support the evolutive conservation of photophore functional organization across luminous etmopterid and dalatiid sharks.