Mammals are among the fastest-radiating groups, being charac terized by a mean species lifespan of the order of 2.5 million years Changes in organisms occur on a variety of temporal scales. Four basic temporal scales (tiers) have been recognized: the ecological timescale; the Milankovitch timescale of precession (the wobbling of the Earth's axis, �21 kyr periodicity), obliquity (tilt of the Earth's axis, 41 kyr periodicity) and eccentricity (the orbit around the Sun, � 100 and �400 kyr periodicity); the million-year timescale of species extinctions and originations; and the ultra-long timescale of mass extinctions and major taxonomic replacements9. Although the main processes controlling the fi rst tier (climate change and competition), the second tier (climate-forced distributional vari ation)3,9,IO and the fourth tier (catastrophic perturbations of the Earth's biosphere) are reasonably well defi ned, the mechanisms underlying third-tier processes are not well ill1derstood. Mammals have featured importantly in discussions on the processes pertaining to this tier2-7• Because their mean species duration is estimated at 2.3-2.6 Myr (refs 1, 2), primary (20-400kyr) Milankovitch vari ations cannot be held responsible for mammal speciation and extinction4,9,lo. It has been suggested that amplitude changes of climatic oscillations could have played a part in explaining turn over3,1O, but until now no efforts have been made systematically to compare such amplitude variations with turnover in long and well dated records.We compiled a data set of more than 200 rodent assemblages from Central Spain (see Supplementary Notes and Supplementary Ta ble 1) and analysed it in terms of turnover. The record is exceptionally long (22 Myr) , largely continuous, dense and well dated. We focused on rodents, because screen sieving allows the collection of large amounts of easily identifi able dental elements. The studied fossils originate from fl uvio-Iacustrine sections in the Madrid, Calatayud-Daroca and Te ruel basins, and amount to over 80,000 isolated molars, which were identifi ed at the species and lineage level (132 lineages, pseudo extinctions ignored). All studied localities are positioned in strati graphic sections, a large number of which have been tied to the geomagnetic polarity timescale by first-order correlations. The complete temporal sequence was calibrated to the new astronomi cally tuned timescale for the Neogene11 (Supplementary Fig. 1). The use of this new timescale is of crucial importance because it allows a direct comparison -with time series of the Earth's orbital parameters outlined above.Randomization procedures were used to capture uncertainties in the ages oflocalities (by the generation of a series of equally probable age models) and of fi rst and last rodent appearances, resulting in 1,000 equally probable time series. Sample size effects on the presence or absence of rodents were observed to be fairly small and were further reduced by inferring lineage presence on a range-through basis and by excluding...

The interest in mammalian palaeohistology has increased dramatically in the last two decades. Starting in 1849 via descriptive approaches, it has been demonstrated that bone tissue and vascularisation types correlate with several biological variables such as ontogenetic stage, growth rate, and ecology. Mammalian bone displays a large variety of bone tissues and vascularisation patterns reaching from lamellar or parallel-fibred to fibrolamellar or woven-fibred bone, depending on taxon and individual age. Here we systematically review the knowledge and methods on cynodont and mammalian bone microstructure as well as palaeohistology and discuss potential future research fields and techniques. We present new data on the bone microstructure of two extant marsupial species and of several extinct continental and island placental mammals. Extant marsupials display mainly parallel-fibred primary bone with radial and oblique but mainly longitudinal vascular canals. Three juvenile specimens of the dwarf island hippopotamid Hippopotamus minor from the Late Pleistocene of Cyprus show reticular to plexiform fibrolamellar bone. The island murid Mikrotia magna from the Late Miocene of Gargano, Italy displays parallel-fibred primary bone with reticular vascularisation and strong remodelling in the middle part of the cortex. Leithia sp., the dormouse from the Pleistocene of Sicily, is characterised by a primary bone cortex consisting of lamellar bone and a high amount of compact coarse cancellous bone. The bone cortex of the fossil continental lagomorph Prolagus oeningensis and three fossil species of insular Prolagus displays mainly parallel-fibred primary bone and reticular, radial as well as longitudinal vascularisation. Typical for large mammals, secondary bone in the giant rhinocerotoid Paraceratherium sp. from the Late Oligocene of Turkey is represented by dense Haversian bone. The skeletochronological features of Sinomegaceros yabei, a large-sized deer from the Pleistocene of Japan closely related to Megaloceros, indicate a high growth rate. These examples and the synthesis of existing data show the potential of bone microstructure to reveal essential information on life history evolution. The bone tissue and the skeletochronological data of the sampled island species suggest the presence of various modes of bone histological modification and mammalian life history evolution on islands to depend on factors of island evolution such as island size, distance from mainland, climate, phylogeny, and time of evolution.

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