E48, these have been tentatively attributed to a time interval that corresponds approximately to the Tortonian/Messinian49. A Late Tortonian (MN11-MN12) or Messinian (MN12-MN13) age represents therefore the best fit for the time of this event of intensified aridification in Gargano and the shift towards a somewhat increased dietary abrasion in Hoplitomeryx. From a wider perspective, this phase of appearance of new LDN193189 cost open-land, arid-adapted vegetation types50 and decreasing humidity51 agrees with the CEP-37440 side effects dominating conditions of the Mediterranean in this epoch. This climatic trend culminated with the Messinian salinity crisis (MN 13, 5.96 Ma), which progressively restricted and finally isolated the Mediterranean Sea from the open ocean52. Evolutionary and ecological implications: island constraints preventing transition among feeding styles. Species of Hoplitomeryx appear to have been sensitive to demographic (high population den-sity), ecological (competition, few resources and food requirements) and abiotic (climate) drivers in Gargano. This variety of causes, probably acting in combination, pushed species to a phase of expansion in diet breadth (i.e., expanding from a soft-leafy to a more abrasive-dominated browsing) preceding strong phenotypic change (e.g., acquisition of extremely hypsodont molar teeth, loss of teeth, evergrowing incisors, shortened premolar series, etc, as recognized in other Mediterranean island ruminants53,54) to escape from overpopulation. Much of the divergence in diet took place during a phase of aridification that favoured the expansion of Hoplitomeryx species into vacant or novel niches. Although a number of additional factors not investigated (such as adjustments in morphology/physiology, geological changes leading to the appearance of novel environments, etc) might influence diversity, diet emerges as paramount in determining ecological diversification on small and resource-limited islands, and represents a density-dependent variable explaining much of the rate and magnitude of insular radiations. It is important to stress, however, that such a dietary expansion in the species did not lead to an immediate change in their major feeding (browsing) type and so, species were not involved in prominent grass-eating. On continents, where mammals adapt more slowly55,56, resources are not limited in variety and extent57 and the diversification dynamics act differently58, the expanded use of different foods among species of Hoplitomeryx may have easily represented the initiation towards a dietary specialization, probably through an initial transition to a more varied diet through a mixed feeding type (i.e., mixture of both browse and grasses), more in accordance with the new environmental circumstances (increased aridity, seasonality and openness of the landscapes) of the epoch. This view is supported by the fact that generalist–both recent and extinct–species are known to better adapt to climatic instability and changing environments than specialized ones40. The following hypothesis needs to be further tested (and the present study implemented through dental microwear in order to offer more specificity and better resolution of the results), but the model here presented strongly supports the view that, despite the potential to exhibit multiple changes in diet composition, the capacityScientific RepoRts | 6:29803 | DOI: 10.1038/srepwww.nature.com/scientificreports/of ruminants to undergo changes in the feeding style on s.E48, these have been tentatively attributed to a time interval that corresponds approximately to the Tortonian/Messinian49. A Late Tortonian (MN11-MN12) or Messinian (MN12-MN13) age represents therefore the best fit for the time of this event of intensified aridification in Gargano and the shift towards a somewhat increased dietary abrasion in Hoplitomeryx. From a wider perspective, this phase of appearance of new open-land, arid-adapted vegetation types50 and decreasing humidity51 agrees with the dominating conditions of the Mediterranean in this epoch. This climatic trend culminated with the Messinian salinity crisis (MN 13, 5.96 Ma), which progressively restricted and finally isolated the Mediterranean Sea from the open ocean52. Evolutionary and ecological implications: island constraints preventing transition among feeding styles. Species of Hoplitomeryx appear to have been sensitive to demographic (high population den-sity), ecological (competition, few resources and food requirements) and abiotic (climate) drivers in Gargano. This variety of causes, probably acting in combination, pushed species to a phase of expansion in diet breadth (i.e., expanding from a soft-leafy to a more abrasive-dominated browsing) preceding strong phenotypic change (e.g., acquisition of extremely hypsodont molar teeth, loss of teeth, evergrowing incisors, shortened premolar series, etc, as recognized in other Mediterranean island ruminants53,54) to escape from overpopulation. Much of the divergence in diet took place during a phase of aridification that favoured the expansion of Hoplitomeryx species into vacant or novel niches. Although a number of additional factors not investigated (such as adjustments in morphology/physiology, geological changes leading to the appearance of novel environments, etc) might influence diversity, diet emerges as paramount in determining ecological diversification on small and resource-limited islands, and represents a density-dependent variable explaining much of the rate and magnitude of insular radiations. It is important to stress, however, that such a dietary expansion in the species did not lead to an immediate change in their major feeding (browsing) type and so, species were not involved in prominent grass-eating. On continents, where mammals adapt more slowly55,56, resources are not limited in variety and extent57 and the diversification dynamics act differently58, the expanded use of different foods among species of Hoplitomeryx may have easily represented the initiation towards a dietary specialization, probably through an initial transition to a more varied diet through a mixed feeding type (i.e., mixture of both browse and grasses), more in accordance with the new environmental circumstances (increased aridity, seasonality and openness of the landscapes) of the epoch. This view is supported by the fact that generalist–both recent and extinct–species are known to better adapt to climatic instability and changing environments than specialized ones40. The following hypothesis needs to be further tested (and the present study implemented through dental microwear in order to offer more specificity and better resolution of the results), but the model here presented strongly supports the view that, despite the potential to exhibit multiple changes in diet composition, the capacityScientific RepoRts | 6:29803 | DOI: 10.1038/srepwww.nature.com/scientificreports/of ruminants to undergo changes in the feeding style on s.
