Although modern beaked whales (Ziphiidae) are known to be highly specialized toothed whales that predominantly feed at great depths upon benthic and benthopelagic prey, only limited palaeontological data document this major ecological shift. We report on a ziphiid-fish assemblage from the Late Miocene of Peru that we interpret as the first direct evidence of a predator-prey relationship between a ziphiid and epipelagic fish. Preserved in a dolomite concretion, a skeleton of the stem ziphiid Messapicetus gregarius was discovered together with numerous skeletons of a clupeiform fish closely related to the epipelagic extant Pacific sardine (Sardinops sagax). Based on the position of fish individuals along the head and chest regions of the ziphiid, the lack of digestion marks on fish remains and the homogeneous size of individuals, we propose that this assemblage results from the death of the whale (possibly via toxin poisoning) shortly after the capture of prey from a single school. Together with morphological data and the frequent discovery of fossil crown ziphiids in deep-sea deposits, this exceptional record supports the hypothesis that only more derived ziphiids were regular deep divers and that the extinction of epipelagic forms may coincide with the radiation of true dolphins.

A new genus and species of Ziphiidae, Dagonodum mojnum gen. nov., sp. nov., from the upper Miocene Gram Formation (c. 9.9-7.2 Ma) represents the first occurrence of the family in Denmark. This long-snouted ziphiid is characterized by two pairs of mandibular tusks, the Eustachian outlet that approximately levels with the dorsalmost margin of the posterior portion of the involucrum, and the left trapezoid nasal with a posteromedial projection into the frontal. A phylogenetic analysis including 25 species and 69 characters was conducted. Dagonodum mojnum is placed in a basal ziphiid clade as the sister taxon of Messapicetus. The specimen is probably a male, because it has enlarged tusks. Alternatively, females could also be involved in fights and develop erupted tusks as in the extant Berardius. Although less well supported, this interpretation proposes that aggressive interactions were not restricted to males in stem-ziphiids. With a thickened thyrohyal and the presence of a precoronoid crest, D. mojnum was able to use suction feeding, but was less specialized to it compared to extant ziphiids. The elongated neck of D. mojnum less optimized to perform deep dives, and the shallow depth at which the Gram Formation was deposited corroborates the hypothesis that at least part of the stem-ziphiids were not regular deep divers.

Abstract. Ziphiidae (beaked whales) are a successful family of medium-to large-sized toothed whales. Their extant members perform regular deep dives beyond the photic zone to forage for cephalopods and fish. Conversely, extinct longsnouted stem ziphiids are interpreted as epipelagic predators. However, some aspects of this hypothesis remain unclear due to the lack of clear morphological proxies for recognizing regular deep divers.We compared the forelimb, neck, and pterygoid sinus system of the fossil ziphiid Messapicetus gregarius with those of other odontocetes to evaluate the potential of these body regions as proxies to assess deep-diving specialization. The reconstructed musculature of the neck and forelimb of M. gregarius was also compared with that of other odontocetes. We also quantified variation in the proportions of the forelimb and the hamular fossa of the pterygoid sinus (HF) using 16 linear measurements. The degree of association between diving behaviour in extant odontocetes and these measurements was evaluated with and without phylogenetic correction.Reconstruction of the neck musculature suggests that M. gregarius possessed a neck more flexible than most extant ziphiids due to the lower degree of fusion of the cervical vertebrae and the large insertions for the M. longus colli and Mm. intertransversarii ventrales cervicis. While neck rigidity might be related to deep diving, differences in neck flexibility among extant ziphiids indicate a more complex functional interpretation. The relationship between forelimb morphology and diving behaviour was not significant, both with and without phylogenetic correction, suggesting that it cannot be used to assess deep-diving abilities with the parameters considered here. Measurements of the HF revealed successful to evaluate deep-diving abilities in odontocetes, with an enlargement of this structure in deep divers. Considering other evidence that suggests an epipelagic behaviour, we propose different scenarios to explain the observation of an enlarged HF in M. gregarius: (1) this species may have fed at different depths; (2) it performed deep dives to avoid potential predators; or (3) the enlarged HF and deep-diving habitat correspond to an ancestral condition, with M. gregarius returning to a more epipelagic habitat.