Pasquale Raia

THE ISLAND RULE IN LARGE MAMMALS: PALEONTOLOGY MEETS ECOLOGY

Abstract The island rule is the phenomenon of the miniaturization of large animals and the gigantism of small animals on islands, with mammals providing the classic case studies. Several explanations for this pattern have been suggested, and departures from the predictions of this rule are common among mammals of differing body size, trophic habits, and phylogenetic affinities. Here we offer a new explanation for the evolution of body size of large insular mammals, using evidence from both living and fossil island faunal assemblages. We demonstrate that the extent of dwarfism in ungulates depends on the existence of competitors and, to a lesser extent, on the presence of predators. In contrast, competition and predation have little or no effect on insular carnivore body size, which is influenced by the nature of the resource base. We suggest dwarfism in large herbivores is an outcome of the fitness increase resulting from the acceleration of reproduction in low‐mortality environments. Carnivore size is dependent on the abundance and size of their prey. Size evolution of large mammals in different trophic levels has different underlying mechanisms, resulting in different patterns. Absolute body size may be only an indirect predictor of size evolution, with ecological interactions playing a major role.

The fast life of a dwarfed giant

In the first half of the 1960s, a rich paleontological site was discovered at Spinagallo caves (Eastern Sicily, Southern Italy). A very abundant fossil population (at least 104 specimens) of the dwarf elephant Elephas falconeri, the smallest elephant that ever lived, was recovered. We computed the survivorship curve for this fossil population in order to investigate both the great juvenile abundance and high calf mortality which it shows. Through the analysis of E. falconeri survivorship, of some reconstructed life-history traits, and of its ecology, and taking into account the Island rule (Foster, 1964), we concluded that E. falconeri moved somewhat toward the ‘fast’ extreme of the slow-fast continuum in life-history traits in regards to its mainland ancestor E. antiquus, that is, it was somehow r-selected. In keeping with our findings, we propose a new explanation for the common occurrence of dwarfism in large mammals living on islands. We suggest the interplay of competition, resource allocation shift and feeding niche width could successfully explain this pattern.

The blue lizard spandrel and the island syndrome

BackgroundMany small vertebrates on islands grow larger, mature later, lay smaller clutches/litters, and are less sexually dimorphic and aggressive than their mainland relatives. This set of observations is referred to as the Island Syndrome. The syndrome is linked to high population density on islands. We predicted that when population density is low and/or fluctuating insular vertebrates may evolve correlated trait shifts running opposite to the Island Syndrome, which we collectively refer to as the reversed island syndrome (RIS) hypothesis. On the proximate level, we hypothesized that RIS is caused by increased activity levels in melanocortin receptors. Melanocortins are postranslational products of the proopiomelanocortin gene, which controls pleiotropically pigmentation, aggressiveness, sexual activity, and food intake in vertebrates.ResultsWe tested the RIS hypothesis performing a number of behavioral, genetic, and ontogenetic tests on a blue colored insular variant of the Italian Wall lizard Podarcis sicula, living on a small island off the Southern Italian coast. The population density of this blue-colored variant was generally low and highly fluctuating from one year to the next.In keeping with our predictions, insular lizards were more aggressive and sexually dimorphic than their mainland relatives. Insular males had wide, peramorphic heads. The growth rate of insular females was slower than growth rates of mainland individuals of both sexes, and of insular males. Consequently, size and shape dimorphism are higher on the Island. As predicted, melanocortin receptors were much more active in individuals of the insular population. Insular lizards have a higher food intake rate than mainland individuals, which is consistent with the increased activity of melanocortin receptors. This may be adaptive in an unpredictable environment such as Licosa Island. Insular lizards of both sexes spent less time basking than their mainland relatives. We suspect this is a by-product (spandrel) of the positive selection for increased activity of melanocortins receptors.ConclusionsWe contend that when population density is either low or fluctuating annually as a result of environmental unpredictability, it may be advantageous to individuals to behave more aggressively, to raise their rate of food intake, and allocate more energy into reproduction.

Ecological specialization in fossil mammals explains Cope's rule.

Cope’s rule is the trend toward increasing body size in a lineage over geological time. The rule has been explained either as passive diffusion away from a small initial body size or as an active trend upheld by the ecological and evolutionary advantages that large body size confers. An explicit and phylogenetically informed analysis of body size evolution in Cenozoic mammals shows that body size increases significantly in most inclusive clades. This increase occurs through temporal substitution of incumbent species by larger-sized close relatives within the clades. These late-appearing species have smaller spatial and temporal ranges and are rarer than the incumbents they replace, traits that are typical of ecological specialists. Cope’s rule, accordingly, appears to derive mainly from increasing ecological specialization and clade-level niche expansion rather than from active selection for larger size. However, overlain on a net trend toward average size increase, significant pulses in origination of large-sized species are concentrated in periods of global cooling. These pulses plausibly record direct selection for larger body size according to Bergmann’s rule, which thus appears to be independent of but concomitant with Cope’s.

Human influence on distribution and extinctions of the late Pleistocene Eurasian megafauna

Late Pleistocene extinctions are of interest to paleontological and anthropological research. In North America and Australia, human occupation occurred during a short period of time and overexploitation may have led to the extinction of mammalian megafauna. In northern Eurasia megafaunal extinctions are believed to have occurred over a relatively longer period of time, perhaps as a result of changing environmental conditions, but the picture is much less clear. To consider megafaunal extinction in Eurasia, we compare differences in the geographical distribution and commonness of extinct and extant species between paleontological and archaeological localities from the late middle Pleistocene to Holocene. Purely paleontological localities, as well as most extinct species, were distributed north of archaeological sites and of the extant species, suggesting that apart from possible differences in adaptations between humans and other species, humans could also have a detrimental effect on large mammal distribution. However, evidence for human overexploitation applies only to the extinct steppe bison Bison priscus. Other human-preferred species survive into the Holocene, including Rangifer tarandus, Equus ferus, Capreolus capreolus, Cervus elaphus, Equus hemionus, Saiga tatarica, and Sus scrofa. Mammuthus primigenius and Megaloceros giganteus were rare in archaeological sites. Carnivores appear little influenced by human presence, although they become rarer in Holocene archaeological sites. Overall, the data are consistent with the conclusion that humans acted as efficient hunters selecting for the most abundant species. Our study supports the idea that the late Pleistocene extinctions were environmentally driven by climatic changes that triggered habitat fragmentation, species range reduction, and population decrease, after which human interference either by direct hunting or via indirect activities probably became critical.

Habitat tracking, stasis and survival in Neogene large mammals

Species response to environmental change may vary from adaptation to the new conditions, to dispersal towards territories with better ecological settings (known as habitat tracking), and to extinction. A phylogenetically explicit analysis of habitat tracking in Caenozoic large mammals shows that species moving over longer distances during their existence survived longer. By partitioning the fossil record into equal time intervals, we showed that the longest distance was preferentially covered just before extinction. This supports the idea that habitat tracking is a key reaction to environmental change, and confirms that tracking causally prolongs species survival. Species covering longer distances also have morphologically less variable cheek teeth. Given the tight relationship between cheek teeth form and habitat selection in large mammals, this supports the well-known, yet little tested, idea that habitat tracking bolsters morphological stasis.

Testing evolutionary stasis and trends in first lower molar shape of extinct Italian populations of Terricola savii (Arvicolidae, Rodentia) by means of geometric morphometrics

Extinct populations of Terricola savii have been investigated in order to analyse evolutionary stasis and correlation of first lower molar shape with climatic proxies by means of geometric morphometrics. Evolutionary stasis, its recognition and explanation are central topics in evolutionary paleobiology. In this study, tooth shape variation of the arvicolid T. savii has been analysed through time. In addition to explicit multivariate tests of stasis based on landmark and semi‐landmark geometric morphometrics, first lower molar M1 shape has been decomposed in orthogonal axes of variation and tested for correlation with climate changes. Multivariate tests were consistent with evolutionary stasis. Yet, according to univariate tests, the dominant dimension of shape variation shows a temporal trend well correlated with a climatic proxy, i.e. δ18O. The remaining variation does not show any trend. Adaptation to current climatic condition might occur even without affecting shape as a whole. Phenotypic plasticity of this species could be invoked to explain evolutionary stasis, as a long time pattern.

Analysis of Heavy Metal Sources for Urban Creeks in the Czech Republic

The study analyzes the presence and the origin of heavy metals in environmental compartments affected by anthropogenic activities. The paper presents the results of a field study performed on the sediments of two of the main small urban creeks of the city of Prague (Czech Republic). The aim of the survey was to verify the presence and bioavailability of heavy metals (Cu, Cr, Ni, Pb, Zn) in the aquatic environment (water as well as bottom sediments), and to assess the source of these pollutants. The results were processed to evaluate the enrichment factor and the partition coefficient, and were statistically analyzed through the analysis of variance and the principal component analysis. Comparison with relevant environmental quality standards showed that measured heavy metal concentrations were always lower than the probable effect concentration (PEC). On the contrary, the threshold effect concentration (TEC) was frequently exceeded. Sequential extraction analysis showed that the bioavailability of studied metals is quite high, suggesting that they could be easily released from the sediment to the aquatic environment. Overall, several sources of pollution, different for the different metals, were identified, all related with anthropogenic factors.

Bioerosion on Brachiopod Shells of a Thanatocoenosis of Alboràn Sea (Spain)

Bioerosion on shells of a brachiopod-bearing thanatocoenosis dredged on red-coral sea-bottom (depth 100 m, 12 miles off the southwestern coast of Alborán Island, Spain) was analyzed. Although low, the intensity of predation was noticeable. Predation drill holes belonging to the ichnogenera Oichnus simplex, O. paraboloides and O. ovalis were recognized. They were probably produced by Muricidae and Marginellidae (gastropods) and octopod cephalopods, respectively. Oichnus ovalis was more common on Terebratulina retusa. A strong selectivity with respect to species, valve, and size was noted. Terebratulina retusa, especially small individuals, was preyed upon more often than any other species. The central portion of both valves was attacked more frequently than expected by chance.

A Dysfunctional Sense of Smell: The Irreversibility of Olfactory Evolution in Free-Living Pigs

Artificial selection began to override natural selection in domesticated wild boar and other species about 10,000xa0years ago. The intentional selection of a desired phenotypic trait is a complex process, and comes along with unexpected or even unwanted changes in other traits, because of epistatic gene effects, and ontogenetic constraints. The loss of brain mass in domestic ungulates is related to selection for reduced reaction to external stimuli. Evolutionary losses in body structures and genes were once considered mostly irreversible, in keeping with Dollo’s law. Here we studied the biochemical and the histological functioning of the free-living pigs (FLPs) olfactory system, to see if and to what extent does FLPs regain a full sense of smell, as compared to the domestic pigs and wild boar Sus scrofa. In our samples both wild boars and FLPs have significantly larger brain per unit mass than domestic pigs, and FLPs’ brains are not significantly smaller than wild boar’s. Similarly, both wild boars and FLPs have significantly higher cell density than domestic pigs in the olfactory mucosa. Yet, at the functional level, olfactory marker protein and neuropeptide Y, both of which are important to the correct functioning of the sense of smell, are fully expressed only in wild boar. These results suggest that FLPs reacquired structural, but not the biochemical capability in their olfactory system.