Virtual reconstruction of the Canis arnensis type (Canidae, Mammalia) from the Upper Valdarno Basin (Italy, Early Pleistocene).

Taphonomic deformation, whether it be brittle or plastic, is possibly the most influential process hindering the correct understanding of fossil species morphology. This is especially true if the deformation affects type specimens or applies to or obscures taxonomically diagnostic or functionally significant traits. Target Deformation, a recently developed virtual manipulation protocol, was implemented to address this issue by applying landmark-guided restoration of the original, deformed fossils, using undeformed specimens (or parts thereof) of the same species as a reference. The enigmatic Early Pleistocene canid Canis arnensis provides a typical example of a fossil species in dire need of virtual restoration. Its lectotype specimen is heavily deformed and none of the few known skulls are well preserved, obscuring the recognition of its systematic and phylogenetic position. Our results indicate that the algorithm effectively countered the lectotype skull's laterolateral compression and its concomitant rostrocaudal elongation. Morphometrically, comparison of the retrodeformed cranium (IGF 867_W) with other specimens of the same species, and to other fossil and extant canid material, confirms IGF 867_W consistently clusters within C. arnensis variability. Overall, the evidence presented here confirms that Target Deformation provides a powerful tool to better characterize complex taxa like C. arnensis, whose knowledge is severely affected by the state of preservation of its fossil material.

Variation in the strength of allometry drives rates of evolution in primate brain shape.

Large brains are a defining feature of primates, as is a clear allometric trend between body mass and brain size. However, important questions on the macroevolution of brain shape in primates remain unanswered. Here we address two: (i), does the relationship between the brain size and its shape follow allometric trends and (ii), is this relationship consistent over evolutionary time? We employ three-dimensional geometric morphometrics and phylogenetic comparative methods to answer these questions, based on a large sample representing 151 species and most primate families. We found two distinct trends regarding the relationship between brain shape and brain size. Hominoidea and Cercopithecinae showed significant evolutionary allometry, whereas no allometric trends were discernible for Strepsirrhini, Colobinae or Platyrrhini. Furthermore, we found that in the taxa characterized by significant allometry, brain shape evolution accelerated, whereas for taxa in which such allometry was absent, the evolution of brain shape decelerated. We conclude that although primates in general are typically described as large-brained, strong allometric effects on brain shape are largely confined to the order's representatives that display more complex behavioural repertoires.

Simultaneous detection of macroevolutionary patterns in phenotypic means and rate of change with and within phylogenetic trees including extinct species.

Recognizing evolutionary trends in phenotypic means and rates requires the application of phylogenetic comparative methods (PCMs). Most PCMs are unsuited to make full use of fossil information, which is a drawback, given the inclusion of such data improves, and in some cases even corrects, the proper understanding of trait evolution. Here we present a new computer application, written in R, that allows the simultaneous computation of temporal trends in phenotypic mean and evolutionary rate along a phylogeny, and to contrast such patterns among different clades within the tree. By using simulation experiments, we show the new implementation, names search.trend is as powerful as existing PCM tools in discerning macroevolutionary patterns in phenotypic means and rates, but differently from any other PCM allows comparing individual clades to each other, and provides rich information about trait evolution for all lineages in the tree.

Unexpectedly rapid evolution of mandibular shape in hominins.

Members of the hominins - namely the so-called 'australopiths' and the species of the genus Homo - are known to possess short and deep mandibles and relatively small incisors and canines. It is commonly assumed that this suite of traits evolved in early members of the clade in response to changing environmental conditions and increased consumption of though food items. With the emergence of Homo, the functional meaning of mandible shape variation is thought to have been weakened by technological advancements and (later) by the control over fire. In contrast to this expectation, we found that mandible shape evolution in hominins is exceptionally rapid as compared to any other primate clade, and that the direction and rate of shape change (from the ape ancestor) are no different between the australopiths and Homo. We deem several factors including the loss of honing complex, canine reduction, and the acquisition of different diets may have concurred in producing such surprisingly high evolutionary rates. This study reveals the evolution of mandibular shape in hominins has strong morpho-functional and ecological significance attached.

Occurrence of enteropathogenic bacteria in birds of prey in Italy.

The importance of wild birds as potential vectors of disease has received recent renewed empirical interest, especially regarding human health although information regarding the enteropathogenic bacteria in birds of prey continue to be scant. This study was performed with the aim to evaluate the occurrence of enteropathogenic bacteria (i.e. Campylobacter spp. Escherichia coli, Salmonella spp.) in birds of prey carcasses in Southern Italy. The results of the present study showed a prevalence of 33·1% (49/148) for Campylobacter spp. where all positive isolates (49/49) were identified as Campylobacter jejuni, and among these positive 12/49 were also identified as Campylobacter coli. Thus, 12/49 birds of prey showed mixed infections for both Campylobacter species. Differences in Campylobacter spp. prevalence between diurnal and nocturnal birds were statistically significant (P = 0·016). Escherichia coli showed a prevalence of 6·8% (10/148) and were serogrouped as O26 (n = 3), O55 (n = 2), O145 (n = 5). Salmonella spp. showed a prevalence of 6·8% (10/148) and were serotyped as S. Napoli (n = 4), Salmonella salamae (n = 3) and S. Typhimurium (n = 3). Although wildlife disease outbreaks have often been underreported in the broader context of global epidemiology, results of the present study suggest that birds of prey may serve as a reservoir of pathogens for livestock and human health, acting at the animal-human-ecosystem interface. SIGNIFICANCE AND IMPACT OF THE STUDY: This study confirms the role of birds of prey as a reservoir of enteropathogenic bacteria (i.e. Campylobacter spp., Escherichia coli, Salmonella spp.). Wild birds can contaminate environment with their faeces and play a crucial role in the transmission of pathogens to poultry and livestock farms and aquifers supplying water to humans. Furthermore, wild birds could disseminate pathogens within rescue and rehabilitation centres where they are admitted.

Like a pig out of water: seaborne spread of domestic pigs in Southern Italy and Sardinia during the Bronze and Iron Ages.

Southern Italy has a long history of human occupation and passage of different cultures since the Early Holocene. Repeated, ancient introductions of pigs in several geographic areas in Europe make it difficult to understand pig translocation and domestication in Italy. The archeozoological record may provide fundamental information on this, hence shedding light on peopling and on trading among different ancient cultures in the Mediterranean. Yet, because of the scanty nature of the fossil record, ancient remains from human-associated animals are somewhat rare. Fortunately, ancient DNA analysis as applied to domestic species proved to be a powerful tool in revealing human migrations. Herein, we analyzed 80-bp fragment of mitochondrial DNA control region from 27 Sus scrofa ancient samples retrieved from Southern Italian and Sardinian archeological sites, spanning in age from the Mesolithic to the Roman period. Our results surprisingly indicate the presence of the Near Eastern haplotype Y1 on both Italy's major islands (Sardinia and Sicily) during the Bronze Age, suggesting the seaborne transportation of domestic pigs by humans at least during 1600-1300 BC. The presence of the Italian E2 clade in domestic contexts shows that the indigenous wild boar was effectively domesticated or incorporated into domestic stocks in Southern Italy during the Bronze Age, although the E2 haplotype has never been found in modern domestic breeds. Pigs belonging to the endemic E2 clade were thus traded between the Peninsula and Sardinia by the end of the second millennium BC and this genetic signature is still detected in Sardinian feral pigs.

Progress to extinction: increased specialisation causes the demise of animal clades.

Animal clades tend to follow a predictable path of waxing and waning during their existence, regardless of their total species richness or geographic coverage. Clades begin small and undifferentiated, then expand to a peak in diversity and range, only to shift into a rarely broken decline towards extinction. While this trajectory is now well documented and broadly recognised, the reasons underlying it remain obscure. In particular, it is unknown why clade extinction is universal and occurs with such surprising regularity. Current explanations for paleontological extinctions call on the growing costs of biological interactions, geological accidents, evolutionary traps, and mass extinctions. While these are effective causes of extinction, they mainly apply to species, not clades. Although mass extinctions is the undeniable cause for the demise of a sizeable number of major taxa, we show here that clades escaping them go extinct because of the widespread tendency of evolution to produce increasingly specialised, sympatric, and geographically restricted species over time.

Venturing out safely: The biogeography of Homo erectus dispersal out of Africa.

The dispersal of Homo erectus out of Africa at some 1.9 million years ago is one of the most important, crucial, and yet controversial events in human evolution. Current opinions about this episode expose the contrast between those who see H. erectus as a highly social, cooperative species seeking out new ecological opportunities to exploit, and those preferring a passive, climate driven explanation for such an event. By using geostatistics techniques and probabilistic models, we characterised the ecological context of H. erectus dispersal, from its East African origin to the colonization of Eurasia, taking into account both the presence of other large mammals and the physical characteristics of the landscape as potential factors. Our model indicated that H. erectus followed almost passively the large herbivore fauna during its dispersal. In Africa, the dispersal was statistically associated with the presence of large freshwater bodies (Rift Valley Lakes). In Eurasia, the presence of H. erectus was associated with the occurrence of geological outcrops likely yielding unconsolidated flint. During the early phase of dispersal, our model indicated that H. erectus actively avoided areas densely populated by large carnivores. This pattern weakened as H. erectus dispersed over Europe, possibly because of the decreasing presence of carnivores there plus the later acquisition of Acheulean technology. During this later phase, H. erectus was associated with limestone and shaley marl, and seems to have been selecting for high-elevation sites. While our results do not directly contradict the idea that H. erectus may have been an active hunter, they clearly point to the fact that predator avoidance may have conditioned its long-distance diffusion as it moved outside Africa. The modelled dispersal route suggests that H. erectus remained preferentially associated with low/middle latitude (i.e., comparatively warm) sites throughout its colonization history.

The conceptual framework of ontogenetic trajectories: parallel transport allows the recognition and visualization of pure deformation patterns.

Ontogeny is usually studied by analyzing a deformation series spanning over juvenile to adult shapes. In geometric morphometrics, this approach implies applying generalized Procrustes analysis coupled with principal component analysis on multiple individuals or multiple species datasets. The trouble with such a procedure is that it mixes intra- and inter-group variation. While MANCOVA models are relevant statistical/mathematical tools to draw inferences about the similarities of trajectories, if one wants to observe and interpret the morphological deformation alone by filtering inter-group variability, a particular tool, namely parallel transport, is necessary. In the context of ontogenetic trajectories, one should firstly perform separate multivariate regressions between shape and size, using regression predictions to estimate within-group deformations relative to the smallest individuals. These deformations are then applied to a common reference (the mean of per-group smallest individuals). The estimation of deformations can be performed on the Riemannian manifold by using sophisticated connection metrics. Nevertheless, parallel transport can be effectively achieved by estimating deformations in the Euclidean space via ordinary Procrustes analysis. This approach proved very useful in comparing ontogenetic trajectories of species presenting large morphological differences at early developmental stages.

Rapid action in the Palaeogene, the relationship between phenotypic and taxonomic diversification in Coenozoic mammals.

A classic question in evolutionary biology concerns the tempo and mode of lineage evolution. Considered variously in relation to resource utilization, intrinsic constraints or hierarchic level, the question of how evolutionary change occurs in general has continued to draw the attention of the field for over a century and a half. Here we use the largest species-level phylogeny of Coenozoic fossil mammals (1031 species) ever assembled and their body size estimates, to show that body size and taxonomic diversification rates declined from the origin of placentals towards the present, and very probably correlate to each other. These findings suggest that morphological and taxic diversifications of mammals occurred hierarchically, with major shifts in body size coinciding with the birth of large clades, followed by taxonomic diversification within these newly formed clades. As the clades expanded, rates of taxonomic diversification proceeded independently of phenotypic evolution. Such a dynamic is consistent with the idea, central to the Modern Synthesis, that mammals radiated adaptively, with the filling of adaptive zones following the radiation.

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.

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.

Longer in the tooth, shorter in the record? The evolutionary correlates of hypsodonty in Neogene ruminants.

The acquisition of hypsodont molars is often regarded as a key innovation in the history of ruminant ungulates. Hypsodont ruminants diversified rapidly during the later Neogene, circa 15-2 Myr ago, and came to dominate the ruminant fossil record in terms of species diversity. Here we show that hypsodont clades had higher speciation and diversification rates than other clades. Hypsodont species had, on average, shorter stratigraphic durations, smaller range size and lower occupancy than non-hypsodont species. Within hypsodont clades, some species were very common and acquired large geographical ranges, whereas others were quite rare and geographically limited. We argue that hypsodont clades diversified in an adaptive radiation-like fashion, with species often splitting cladogenetically while still in the expansive phase of their occupancy history.

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 M(1) 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. delta(18)O. 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.

Evaluation of cardiac and respiratory involvement in sarcoglycanopathies.

Sarcoglycanopathies constitute a subgroup of limb-girdle recessive muscular dystrophies due to defects in sarcoglycan complex that comprises five distinct transmembrane proteins called alpha-, beta-, gamma-, delta-and epsilon-sarcoglycans. As it is well known that sarcoglycans are expressed both in heart and in skeletal muscles and a complete deficiency in delta-sarcoglycan is the cause of the Syrian hamster BIO.14 cardiomyopathy, we studied cardiac and respiratory involvement in 20 patients with sarcoglycanopathies by clinical, electrocardiographic, echocardiographic, scintigraphic and spirometric assessments. A normal heart function was found in 31.3% of all patients; a preclinical cardiomyopathy in 43.7%; an arrhythmogenic cardiomyopathy in 6.3% and initial signs of dilated cardiomyopathy in 18.7%. In one patient the data were examined retrospectively. No correlation was found between cardiac and skeletal muscle involvement. With reference to the type of sarcoglycanopathy, signs of hypoxic myocardial damage occurred in beta-, gamma- and delta-sarcoglycanopathies, while initial signs of a dilated cardiomyopathy in gamma- and delta-sarcoglycanopathies were found. A normal respiratory function was observed in 23.5% of all patients, a mild impairment in 35.4%, a moderate impairment in 29.4%, and a severe impairment in 11.7%.

Left ventricular function and perfusion in Becker's muscular dystrophy.

UNLABELLED: The aim of this study was to evaluate left ventricular (LV) perfusion and function in patients with Becker muscular dystrophy (BMD). METHODS: Fourteen male patients (age range 14-40 yr) with BMD were evaluated by 201Tl SPECT and radionuclide angiography both at rest and after dipyridamole stress test. RESULTS: All patients showed uptake defect demonstrated by 201Tl SPECT (mean 4.1 +/- 2.2 uptake defect/patient). Significant relationships (p < 0.05) were found between the number of uptake defects and rest LV ejection fraction (LVEF) (r = -0.54); peak filling rate (PFR) (r = -0.57) and dipyridamole LVEF (r = -0.65). Dipyridamole induced reversible uptake defects were found in 7/14 (50%) patients with BMD. The 14 patients were divided into two groups on the basis of the presence (Group A, n = 6) or the absence (Group B, n = 8) of severe irreversible uptake defect (i.e., < 50% 201Tl uptake). Group A showed lower values of PFR and LVEF when compared to patients of Group B. CONCLUSIONS: In patients with BMD there is a relatively high incidence of uptake defects and LV function (both at rest and after dipyridamole) appears to be related to the number of uptake defects. Moreover, the presence of severe irreversible uptake defects identifies a subgroup of patients with BMD characterized by a severely depressed LV function.

Systolic and diastolic interaction in the assessment of left ventricular function following surgical cardioplegia.

Using regionally implanted sonomicrometry crystals, we have evaluated a new index of regional function (normalised systolic shortening [NSS]) which integrates the systolic and diastolic properties of the left ventricle. Eight dogs (group I) were subjected to standard cardiopulmonary bypass and 45 min of hypothermic (10 degrees C), hyperkalaemic (25 mEq) crystalloid cardioplegia. Seven dogs (group II) underwent occlusion of the left anterior descending (LAD) coronary artery 5 min prior to initiation of cardiac arrest. The occlusion was released after 20 min, before the second cardioplegia infusion. Sarnoff left ventricular (LV) function curves were performed pre-arrest and 20, 40 and 60 min after removal of the cross-clamp. Regional assessment of myocardial function showed 55% +/- 3%, 70% +/- 3% and 70% +/- 5% recovery in the LAD region and 52% +/- 2%, 83% +/- 3% and 88% +/- 4% recovery in the circumflex (Cx) region of group I. In group II the LAD region recovered 27% +/- 1%, 31% +/- 3% and 38% +/- 3% and the Cx region showed 61% +/- 3%, 55% +/- 1% and 65% +/- 5% recovery. Comparison of the new index of ventricular function to standard indices of regional and global function demonstrate that the latter underestimate the degree of myocardial dysfunction after cardioplegic arrest, particularly in situations of acute regional myocardial ischaemia and uneven myocardial protection. The utilization of this index should provide a better standard for the more accurate assessment of interventions designed to decrease myocardial injury during cardioplegic arrest.