Evolution of morphological adaptations for digging in living and extinct ctenomyid and octodontid rodents

To examine the evolution of burrowing specializations in the sister families Octodontidae and Ctenomyidae (Rodentia: Caviomorpha), we produced a synthetic phylogeny (supertree), combining both molecular and morphological phylogenies, and including both fossil and extant genera. We mapped morphological specializations of the digging apparatus onto our phylogenetic hypothesis and attempted to match morphological diversity with information on the ecology and behaviour of octodontoid taxa. Burrowing for sheltering and rearing is the rule among octodontids and ctenomyids, and adaptations for digging have been known from the Early Pliocene onward. However, only a few taxa have evolved fully subterranean habits. Scratch-digging is widespread among both semifossorial and fully subterranean lineages, and morphological changes associated with scratch-digging are not restricted to subterranean lineages. By contrast, various adaptations for chisel-tooth digging are restricted to some subterranean lineages and are combined differently in the octodontid Spalacopus, the fossil ctenomyid Eucelophorus, and some living Ctenomys. Some octodontid taxa are able to dig complex burrows in spite of having no substantial changes in musculoskeletal attributes. Hence, we suggest that, during the early evolution of those branches giving rise to fully subterranean ctenomyids and octodontids, a change in behaviour probably preceded the origin of structural adaptations.Facultad de Ciencias Naturales y Muse

Evolution of morphological adaptations for digging in living and extinct ctenomyid and octodontid rodents

To examine the evolution of burrowing specializations in the sister families Octodontidae and Ctenomyidae (Rodentia: Caviomorpha), we produced a synthetic phylogeny (supertree), combining both molecular and morphological phylogenies, and including both fossil and extant genera. We mapped morphological specializations of the digging apparatus onto our phylogenetic hypothesis and attempted to match morphological diversity with information on the ecology and behaviour of octodontoid taxa. Burrowing for sheltering and rearing is the rule among octodontids and ctenomyids, and adaptations for digging have been known from the Early Pliocene onward. However, only a few taxa have evolved fully subterranean habits. Scratch-digging is widespread among both semifossorial and fully subterranean lineages, and morphological changes associated with scratch-digging are not restricted to subterranean lineages. By contrast, various adaptations for chisel-tooth digging are restricted to some subterranean lineages and are combined differently in the octodontid Spalacopus, the fossil ctenomyid Eucelophorus, and some living Ctenomys. Some octodontid taxa are able to dig complex burrows in spite of having no substantial changes in musculoskeletal attributes. Hence, we suggest that, during the early evolution of those branches giving rise to fully subterranean ctenomyids and octodontids, a change in behaviour probably preceded the origin of structural adaptations.Facultad de Ciencias Naturales y Muse

Evolution of morphological adaptations for digging in living and extinct ctenomyid and octodontid rodents

To examine the evolution of burrowing specializations in the sister families Octodontidae and Ctenomyidae (Rodentia: Caviomorpha), we produced a synthetic phylogeny (supertree), combining both molecular and morphological phylogenies, and including both fossil and extant genera. We mapped morphological specializations of the digging apparatus onto our phylogenetic hypothesis and attempted to match morphological diversity with information on the ecology and behaviour of octodontoid taxa. Burrowing for sheltering and rearing is the rule among octodontids and ctenomyids, and adaptations for digging have been known from the Early Pliocene onward. However, only a few taxa have evolved fully subterranean habits. Scratch-digging is widespread among both semifossorial and fully subterranean lineages, and morphological changes associated with scratch-digging are not restricted to subterranean lineages. By contrast, various adaptations for chisel-tooth digging are restricted to some subterranean lineages and are combined differently in the octodontid Spalacopus, the fossil ctenomyid Eucelophorus, and some living Ctenomys. Some octodontid taxa are able to dig complex burrows in spite of having no substantial changes in musculoskeletal attributes. Hence, we suggest that, during the early evolution of those branches giving rise to fully subterranean ctenomyids and octodontids, a change in behaviour probably preceded the origin of structural adaptations.Facultad de Ciencias Naturales y Muse

Dispersal and population structure at different spatial scales in the subterranean rodent Ctenomys australis

<p>Abstract</p> <p><b>Background</b></p> <p>The population genetic structure of subterranean rodent species is strongly affected by demographic (e.g. rates of dispersal and social structure) and stochastic factors (e.g. random genetic drift among subpopulations and habitat fragmentation). In particular, gene flow estimates at different spatial scales are essential to understand genetic differentiation among populations of a species living in a highly fragmented landscape. <it>Ctenomys australis </it>(the sand dune tuco-tuco) is a territorial subterranean rodent that inhabits a relatively secure, permanently sealed burrow system, occurring in sand dune habitats on the coastal landscape in the south-east of Buenos Aires province, Argentina. Currently, this habitat is threatened by urban development and forestry and, therefore, the survival of this endemic species is at risk. Here, we assess population genetic structure and patterns of dispersal among individuals of this species at different spatial scales using 8 polymorphic microsatellite loci. Furthermore, we evaluate the relative importance of sex and habitat configuration in modulating the dispersal patterns at these geographical scales.</p> <p>Results</p> <p>Our results show that dispersal in <it>C. australis </it>is not restricted at regional spatial scales (~ 4 km). Assignment tests revealed significant population substructure within the study area, providing support for the presence of two subpopulations from three original sampling sites. Finally, male-biased dispersal was found in the Western side of our study area, but in the Eastern side no apparent philopatric pattern was found, suggesting that in a more continuous habitat males might move longer distances than females.</p> <p>Conclusions</p> <p>Overall, the assignment-based approaches were able to detect population substructure at fine geographical scales. Additionally, the maintenance of a significant genetic structure at regional (~ 4 km) and small (less than 1 km) spatial scales despite apparently moderate to high levels of gene flow between local sampling sites could not be explained simply by the linear distance among them. On the whole, our results support the hypothesis that males disperse more frequently than females; however they do not provide support for strict philopatry within females.</p

Measurement of the cosmic ray spectrum above 4×10184{\times}10^{18} eV using inclined events detected with the Pierre Auger Observatory

A measurement of the cosmic-ray spectrum for energies exceeding $4{\times}10^{18}$ eV is presented, which is based on the analysis of showers with zenith angles greater than $60^{\circ}$ detected with the Pierre Auger Observatory between 1 January 2004 and 31 December 2013. The measured spectrum confirms a flux suppression at the highest energies. Above $5.3{\times}10^{18}$ eV, the "ankle", the flux can be described by a power law $E^{-\gamma}$ with index $\gamma=2.70 \pm 0.02 \,\text{(stat)} \pm 0.1\,\text{(sys)}$ followed by a smooth suppression region. For the energy ($E_\text{s}$) at which the spectral flux has fallen to one-half of its extrapolated value in the absence of suppression, we find $E_\text{s}=(5.12\pm0.25\,\text{(stat)}^{+1.0}_{-1.2}\,\text{(sys)}){\times}10^{19}$ eV.Comment: Replaced with published version. Added journal reference and DO

Energy Estimation of Cosmic Rays with the Engineering Radio Array of the Pierre Auger Observatory

The Auger Engineering Radio Array (AERA) is part of the Pierre Auger Observatory and is used to detect the radio emission of cosmic-ray air showers. These observations are compared to the data of the surface detector stations of the Observatory, which provide well-calibrated information on the cosmic-ray energies and arrival directions. The response of the radio stations in the 30 to 80 MHz regime has been thoroughly calibrated to enable the reconstruction of the incoming electric field. For the latter, the energy deposit per area is determined from the radio pulses at each observer position and is interpolated using a two-dimensional function that takes into account signal asymmetries due to interference between the geomagnetic and charge-excess emission components. The spatial integral over the signal distribution gives a direct measurement of the energy transferred from the primary cosmic ray into radio emission in the AERA frequency range. We measure 15.8 MeV of radiation energy for a 1 EeV air shower arriving perpendicularly to the geomagnetic field. This radiation energy -- corrected for geometrical effects -- is used as a cosmic-ray energy estimator. Performing an absolute energy calibration against the surface-detector information, we observe that this radio-energy estimator scales quadratically with the cosmic-ray energy as expected for coherent emission. We find an energy resolution of the radio reconstruction of 22% for the data set and 17% for a high-quality subset containing only events with at least five radio stations with signal.Comment: Replaced with published version. Added journal reference and DO

Measurement of the Radiation Energy in the Radio Signal of Extensive Air Showers as a Universal Estimator of Cosmic-Ray Energy

We measure the energy emitted by extensive air showers in the form of radio emission in the frequency range from 30 to 80 MHz. Exploiting the accurate energy scale of the Pierre Auger Observatory, we obtain a radiation energy of 15.8 \pm 0.7 (stat) \pm 6.7 (sys) MeV for cosmic rays with an energy of 1 EeV arriving perpendicularly to a geomagnetic field of 0.24 G, scaling quadratically with the cosmic-ray energy. A comparison with predictions from state-of-the-art first-principle calculations shows agreement with our measurement. The radiation energy provides direct access to the calorimetric energy in the electromagnetic cascade of extensive air showers. Comparison with our result thus allows the direct calibration of any cosmic-ray radio detector against the well-established energy scale of the Pierre Auger Observatory.Comment: Replaced with published version. Added journal reference and DOI. Supplemental material in the ancillary file

Factors affecting survival in Mediterranean populations of the Eurasian eagle owl

The survival rate is a key parameter for population management and the monitoring of populations. Thus, an analysis of survival rate variations and the factors influencing the same is essential for understanding population dynamics. Here, we study the factors determining the survival and the causes of mortality of the Eurasian eagle owl (Bubo bubo) in two Spanish Mediterranean populations (Murcia and Seville) where the species has a high population density and breeding success; yet its survival rates and the factors that affect them are unknown. Between 2003 and 2010, 63 breeding owls were captured and radio-tracked. Three monthly (quarterly) survival rates were estimated using known-fate models in the program MARK. The mean overall annual survival rate was 0.776 (95 % CI: 0.677, 0.875). We observed survival differences between sexes, and between the breeding and non-breeding periods, although no overwhelming support was found for any particular model. We concluded that (i) females have a lower survival rate than males, probably due to their larger home ranges, which increase the risk of mortality; (ii) the survival rates of both sexes were lower during the non-breeding period; and (iii) the causes of mortality differed significantly between the two populations, gunshot being the main cause in Seville and electrocution in Murcia.Peer Reviewe