Mikael Fortelius

Approaching a state shift in Earth’s biosphere

Localized ecological systems are known to shift abruptly and irreversibly from one state to another when they are forced across critical thresholds. Here we review evidence that the global ecosystem as a whole can react in the same way and is approaching a planetary-scale critical transition as a result of human influence. The plausibility of a planetary-scale ‘tipping point’ highlights the need to improve biological forecasting by detecting early warning signs of critical transitions on global as well as local scales, and by detecting feedbacks that promote such transitions. It is also necessary to address root causes of how humans are forcing biological changes.

Functional Characterization of Ungulate Molars Using the Abrasion-Attrition Wear Gradient: A New Method for Reconstructing Paleodiets

Abstract The analysis of fossil ungulate cheek teeth has long been one of the main sources of information about the terrestrial environments of the Cenozoic, but the methods used to extract this information have been either imprecise or prohibitively laborious. Here we present a method based on relative facet development that is quantitative, robust, and rapid. This method, which we term mesowear analysis, is based on the physical properties of ungulate foods as reflected in the relative amounts of attritive and abrasive wear that they cause on the dental enamel of the occlusal surfaces. Mesowear was recorded by examining the buccal apices of molar tooth cusps. Apices were characterized as sharp, rounded, or blunt, and the valleys between them either high or low. The method has been developed only for selenodont and trilophodont molars, but the principle is readily extendable to other crown types. Mesowear analysis is insensitive to wear stage as long as the very early and very late stages are excluded. Cluster analysis of the mesowear variables produces clusters reflecting four main groups from abrasion-dominated to attrition-dominated: grazers, graze-dominated mixed feeders, browse-dominated mixed feeders, and browsers. Most of the relatively few apparent anomalies are explained by more detailed dietary information. Mesowear analysis provides resolution within the main dietary classes and the clustering is virtually identical with and without the index of hypsodonty. Discriminant analysis using all mesowear variables and hypsodonty showed an overall correct classification of 76% of 64 species of living ungulates into the conventional dietary categories of browser, grazer, and mixed feeder, while a smaller set of 27 “typical” species was correctly classified at 96%. Alternative “conservative” and “radical” dietary classifications that were employed to accommodate cases where dietary information was controversial or unclear produced only marginally different results. Mesowear analysis successfully resolved a test case using the Serengeti grazing succession and appears to be superior to microwear analysis in two cases where the diet of fossil ungulates has been previously studied by microwear and other conventional methods.

ON THE MEANS WHEREBY MAMMALS ACHIEVE INCREASED FUNCTIONAL DURABILITY OF THEIR DENTITIONS, WITH SPECIAL REFERENCE TO LIMITING FACTORS

( I ) Types of solutions available . . . . . . . . . (2) Increased wear resistance of dental tissues . . . . . . (3) Increased tooth size . . . . . . . . . . . (4) Additional teeth, and a discussion of bilophodonty . . , . , (5) Increased tooth height . . . . . . . . . . (6) Combinations of methods . . . . . . . . . . IV. Conclusion . . . . . . . . . . . . . V. Acknowledgements. . . . . . . . . . . . VI. References . . . . . . . . . . . . . (2) Functional demands, with a note on dental wear

High-level similarity of dentitions in carnivorans and rodents.

The study of mammalian evolution depends greatly on understanding the evolution of teeth and the relationship of tooth shape to diet. Links between gross tooth shape, function and diet have been proposed since antiquity, stretching from Aristotle to Cuvier, Owen and Osborn. So far, however, the possibilities for exhaustive, quantitative comparisons between greatly different tooth shapes have been limited. Cat teeth and mouse teeth, for example, are fundamentally distinct in shape and structure as a result of independent evolutionary change over tens of millions of years. There is difficulty in establishing homology between their tooth components or in summarizing their tooth shapes, yet both carnivorans and rodents possess a comparable spectrum of dietary specializations from animals to plants. Here we introduce homology-free techniques to measure the phenotypic complexity of the three-dimensional shape of tooth crowns. In our geographic information systems (GIS) analysis of 441 teeth from 81 species of carnivorans and rodents, we show that the surface complexity of tooth crowns directly reflects the foods they consume. Moreover, the absolute values of dental complexity for individual dietary classes correspond between carnivorans and rodents, illustrating a high-level similarity between overall tooth shapes despite a lack of low-level similarity of specific tooth components. These results suggest that scale-independent forces have determined the high-level dental shape in lineages that are widely divergent in size, ecology and life history. This link between diet and phenotype will be useful for inferring the ecology of extinct species and illustrates the potential of fast-throughput, high-level analysis of the phenotype.

Molar Tooth Diversity, Disparity, and Ecology in Cenozoic Ungulate Radiations

A classic example of adaptive radiation is the diversification of Cenozoic ungulates into herbivore adaptive zones. Their taxonomic diversification has been associated with changes in molar tooth morphology. Analysis of molar crown types of the Artiodactyla, Perissodactyla, and archaic ungulates (“Condylarthra”) shows that the diversity of genera and crown types was high in the Eocene. Post-Eocene molars of intermediate crown types are rare, and thus the ungulate fauna contained more taxa having fewer but more disparate crown types. Taxonomic diversity trends alone give incomplete descriptions of adaptive radiations.

The Evolution of Maximum Body Size of Terrestrial Mammals

How Mammals Grew in Size Mammals diversified greatly after the end-Cretaceous extinction, which eliminated the dominant land animals (dinosaurs). Smith et al. (p. 1216) examined how the maximum size of mammals increased during their radiation in each continent. Overall, mammal size increased rapidly, then leveled off after about 25 million years. This pattern holds true on most of the continents—even though data are sparse for South America—and implies that mammals grew to fill available niches before other environmental and biological limits took hold. Maximum mammal size increased at the beginning of the Cenozoic, then leveled off after about 25 million years. The extinction of dinosaurs at the Cretaceous/Paleogene (K/Pg) boundary was the seminal event that opened the door for the subsequent diversification of terrestrial mammals. Our compilation of maximum body size at the ordinal level by sub-epoch shows a near-exponential increase after the K/Pg. On each continent, the maximum size of mammals leveled off after 40 million years ago and thereafter remained approximately constant. There was remarkable congruence in the rate, trajectory, and upper limit across continents, orders, and trophic guilds, despite differences in geological and climatic history, turnover of lineages, and ecological variation. Our analysis suggests that although the primary driver for the evolution of giant mammals was diversification to fill ecological niches, environmental temperature and land area may have ultimately constrained the maximum size achieved.

Common mammals drive the evolutionary increase of hypsodonty in the Neogene

During the past 20 million years, herbivorous mammals of numerous lineages have evolved hypsodont, or high-crowned, cheek teeth. Hypsodonty is informative ecologically because it is well developed in mammals eating fibrous and abrasive foods that are most abundant in open and generally or seasonally dry environments. Here we report that in the Neogene of Europe mammals with the greatest locality coverages showed an increase in hypsodonty. We used a data set of 209 localities to measure whether large mammals occurring in many fossil localities show a similar increase in hypsodonty to mammals occurring in single or few localities. Taxonomic and morphological groupings show a low average hypsodonty in the early Miocene epoch. From the middle Miocene onwards, only the hypsodonty of commonly found mammals shows a marked increase. Therefore, in the drying Europe of the late Miocene, only increasingly hypsodont mammals may have been able to expand their share of habitats and food resources. These results suggest that the relatively small number of species known from multiple localities are palaeoecologically informative by themselves, irrespective of the rest of the known species.

Adaptive radiation of multituberculate mammals before the extinction of dinosaurs

The Cretaceous–Paleogene mass extinction approximately 66 million years ago is conventionally thought to have been a turning point in mammalian evolution. Prior to that event and for the first two-thirds of their evolutionary history, mammals were mostly confined to roles as generalized, small-bodied, nocturnal insectivores, presumably under selection pressures from dinosaurs. Release from these pressures, by extinction of non-avian dinosaurs at the Cretaceous–Paleogene boundary, triggered ecological diversification of mammals. Although recent individual fossil discoveries have shown that some mammalian lineages diversified ecologically during the Mesozoic era, comprehensive ecological analyses of mammalian groups crossing the Cretaceous–Paleogene boundary are lacking. Such analyses are needed because diversification analyses of living taxa allow only indirect inferences of past ecosystems. Here we show that in arguably the most evolutionarily successful clade of Mesozoic mammals, the Multituberculata, an adaptive radiation began at least 20 million years before the extinction of non-avian dinosaurs and continued across the Cretaceous–Paleogene boundary. Disparity in dental complexity, which relates to the range of diets, rose sharply in step with generic richness and disparity in body size. Moreover, maximum dental complexity and body size demonstrate an adaptive shift towards increased herbivory. This dietary expansion tracked the ecological rise of angiosperms and suggests that the resources that were available to multituberculates were relatively unaffected by the Cretaceous–Paleogene mass extinction. Taken together, our results indicate that mammals were able to take advantage of new ecological opportunities in the Mesozoic and that at least some of these opportunities persisted through the Cretaceous–Paleogene mass extinction. Similar broad-scale ecomorphological inventories of other radiations may help to constrain the possible causes of mass extinctions.

Distribution history and climatic controls of the Late Miocene Pikermian chronofauna

The Late Miocene development of faunas and environments in western Eurasia is well known, but the climatic and environmental processes that controlled its details are incompletely understood. Here we map the rise and fall of the classic Pikermian fossil mammal chronofauna between 12 and 4.2 Ma, using genus-level faunal similarity between localities. To directly relate land mammal community evolution to environmental change, we use the hypsodonty paleoprecipitation proxy and paleoclimate modeling. The geographic distribution of faunal similarity and paleoprecipitation in successive timeslices shows the development of the open biome that favored the evolution and spread of the open-habitat adapted large mammal lineages. In the climate model run, this corresponds to a decrease in precipitation over its core area south of the Paratethys Sea. The process began in the latest Middle Miocene and climaxed in the medial Late Miocene, about 7–8 million years ago. The geographic range of the Pikermian chronofauna contracted in the latest Miocene, a time of increasing summer drought and regional differentiation of habitats in Eastern Europe and Southwestern Asia. Its demise at the Miocene-Pliocene boundary coincides with an environmental reversal toward increased humidity and forestation, changes inevitably detrimental to open-adapted, wide-ranging large mammals.

Higher origination and extinction rates in larger mammals.

Do large mammals evolve faster than small mammals or vice versa? Because the answer to this question contributes to our understanding of how life-history affects long-term and large-scale evolutionary patterns, and how microevolutionary rates scale-up to macroevolutionary rates, it has received much attention. A satisfactory or consistent answer to this question is lacking, however. Here, we take a fresh look at this problem using a large fossil dataset of mammals from the Neogene of the Old World (NOW). Controlling for sampling biases, calculating per capita origination and extinction rates of boundary-crossers and estimating survival probabilities using capture-mark-recapture (CMR) methods, we found the recurring pattern that large mammal genera and species have higher origination and extinction rates, and therefore shorter durations. This pattern is surprising in the light of molecular studies, which show that smaller animals, with their shorter generation times and higher metabolic rates, have greater absolute rates of evolution. However, higher molecular rates do not necessarily translate to higher taxon rates because both the biotic and physical environments interact with phenotypic variation, in part fueled by mutations, to affect origination and extinction rates. To explain the observed pattern, we propose that the ability to evolve and maintain behavior such as hibernation, torpor and burrowing, collectively termed “sleep-or-hide” (SLOH) behavior, serves as a means of environmental buffering during expected and unexpected environmental change. SLOH behavior is more common in some small mammals, and, as a result, SLOH small mammals contribute to higher average survivorship and lower origination probabilities among small mammals.