Anna Loy

Geometric morphometric analyses of leaf variation in four oak species of the subgenus Quercus (Fagaceae)

Abstract This paper defines a protocol for the study of leaf morphology and morphometric relationships among Quercus frainetto, Quercus robur, Quercus petraea and Quercus pubescens. Leaf shapes were investigated in an attempt to characterize and discriminate among these four species. Landmark and outline data were compared in order to ascertain their relative efficacy for describing variation in leaf morphology. MANOVA was performed to assess intra‐specific differentiation and to compare the distribution of variance in individuals, populations and species. A strong correlation was found between landmark and outline datasets of the four species: both methods proved capable of discriminating among the four species. Finally, a Two‐Block Partial Least‐Squares analysis was performed for each species in order to assess the correlation between intra‐specific shape variation and environmental factors.

Connectivity providers for semi-aquatic vertebrates: the case of the endangered otter in Italy

Modeling habitat connectivity for conservation of semi-aquatic vertebrates is a particularly challenging task, due to the fine-scale and linear distribution of riverine habitats and to the capacity of species to move both on freshwater and terrestrial realms. We showed how the integrated analytical framework provided by the habitat availability (reachability) metrics and their fractions can be used to effectively evaluate the distinctive roles and contributions of both habitat patches (aquatic and riparian) and linkage areas (permeable land matrix) to the connectivity and functioning of a complex system composed of multiple river catchments. Analysis focused on the Eurasian otter (Lutra lutra L.), one of the most endangered mammals in Italy. We developed a network connectivity model based on suitable otter habitats and multiple least-cost paths between catchments. A graph analytical approach was used to identify critical nodes and links for the potential expansion and long-term viability of the species in the region. Our results showed that few basins concentrate most of the importance for sustaining the overall habitat connectivity, due to the extension of suitable habitats they contain, their strong connections with other basins, and their importance as stepping stones that uphold ecological fluxes between otherwise weakly connected habitats. The potential contribution of each basin to enhance the dispersal and expansion of otters in the area strongly depended on the key functional paths (sequences of links and nodes) among the catchments. We identified vacant basins that could be colonized by otters in the near future, and connecting areas in the intermediate matrix that might be preferentially used to conduct and promote dispersal movements and gene flow in the area. The novel approach here adopted could be easily extended to other semiaquatic species and catchment systems, offering a management strategy to preserve the hydrographic network as an integrated system, as well as a joint evaluation of the role of both the river courses and the matrix in between in a single landscape model.

Cranial morphology of Martes foina and M. martes (Mammalia, Carnivora, Mustelidae): The role of size and shape in sexual dimorphism and interspecific differentiation

Abstract Landmark based morphometrics was used to investigate sexual dimorphism and interspecific differentiation in the mustelids Martes foina and M. martes, which are typical sexual size dimorphic species. Analyses were run on 192 specimens of M. foina and 36 specimens of M. martes from central and northern Italy, by recording 19 and 40 landmarks on the dorsal skull and on the palate, respectively. Results indicate that both size and shape are important components of skull dimorphism in males and females of the two species. The skull of the stone marten males is larger, has a narrower postorbital constriction and larger molars than in females. These differences and the partial least square analysis run on shape variable and stomach contents of one sample suggest that stone marten males are better adapted to catch and kill live prey, while females seem to prey upon easier‐to‐catch food. Shape features involved in sexual dimorphism are similar to those involved in interspecific variation, but the two species differ also in the relative size of their canines, carnassial and molar teeth, which are likely related to different ways of acquiring and processing preys.

Genetic and morphometric evidence of introgression between two species of moles (Insectivora: Talpa europaea and Talpa romana ) in central Italy

Allozyme and morphometric variation was studied in two populations of moles, Talpa europaea and T. romana, in an area of sympatric occurrence (Palazzo, near Assisi, central Italy) located along the known parapatric contact boundary. The electrophoretic analysis revealed 36 moles characterized by a genome of T. europaea, and 29 moles with a genome of T. romana. The extension of the overlap zone was c. 2 km along the north–south direction. Fourteen specimens recognized as T. europaea had an Ada allele typical of T. romana, while two specimens identified as T. romana were characterized by two alleles (at the Adh and Sdh loci) previously found only in T. europaea. The skulls of the specimens with introgressed alleles showed features intermediate between those typical of T. romana and T. europaea. However, the intermediate skull shape of the introgressed specimens did not correspond to an intermediate size between T. romana and T. europaea. The occurrence of individuals with introgressed alleles, and the lack of F1 hybrids or backcrossed individuals suggest that at present free interbreeding between the two species is unlikely, but that hybridization and backcrossing has occurred, probably when the species first came into contact.

Territorial behavior in Talpa romana, a fossorial insectivore from southcentral Italy

A natural population of Talpa romana was radiotracked in central Italy, close to the type locality of the species. We assessed the territorial behavior of the species, which is inferred from use of exclusive core areas. Some spatial overlap was observed between the home ranges of animals of different sex, more specifically between the home range of a male and those of neighboring females. Temporal variation of core areas suggests that they represent portions of the home range intensively exploited for food resources on a short-term basis, while maintenance of the entire home range is possibly related to the probability of mating and availability of food in the long term.

Cranial morphometrics of the Apennine brown bear (Ursus arctos marsicanus) and preliminary notes on the relationships with other southern European populations

The geographic variation of seven southern European populations of Ursus arctos was investigated through multivariate morphometrics of the skull. Univariate and multivariate analyses were run on 63 cranial and dental measurements recorded on 50 specimens from the Apennines, the Alps, and the Transcaucasus, and on 14 specimens of the fossil Ursus spelaeus. Detailed analyses of sexual dimorphism, age structure, and ranges of variation have been carried out on the Apennines population, which morphological variation has been little explored. Subsets of characters were selected to allow comparison with data derived from literature for populations from the Pyrenees, the Rhodopi‐Rila‐Pirin, the Balkans, and the Caucasus. Analyses clearly indicate that the Apennine bear is morphologically distinct from both a western (Alps, Pyrenees, Balkans, and Rhodopi) and an eastern contingent (Caucasus and Transcaucasus), therefore suggesting that the Apennine population should be reconsidered as a separate taxon, namely Ursus arctos marsicanus. These preliminary results suggest caution in restocking conservation actions.

Distribution of Talpa europea (Mammalia, Insectivora, Talpidae) in Europe: A biogeographic hypothesis based on morphometric data

Abstract Geographic variation of the European mole Talpa europaea was investigated through univariate and multivariate analyses of mor phometric characters. Fifteen Cartesian coordinates were recorded from the left mandible of 466 specimens from 145 European localities. Sexes and sample localities were pooled after testing for homogeneity of groups. Data were transformed to avoid the effect of within‐group allometry. Four major phenetic clusters are evident, i.e., a French‐Spanish, a central European, an Italian, and a British one. Distribution of specimens in canonical space is related to both geography and climate. The influence of historical events on variation was tested through the analyses of hypothetical secondary contact zones, showing the occurrence of stepclines. These step‐clines may be related to range expansion following ice retreat after the last Pleistocene glaciation.

Morphometric divergence in southern European moles (Insectívora, Talpidae)

Abstract Evolutionary divergence in southern European moles (Talpa eu‐ropaea T. romana, T. stankovici, T. caeca caeca, T. c. hercegoviniensis) was investigated through multivariate morphometrics. The high discrimination shown by taxa seems to be related to geographic independent evolution that occurred during the Pleistocene. Whilst the Balkanic T. c. hercegoviniensis is morphometrically closest to T. caeca caeca, slight chromosomal diversity suggests active speciation.

Digging adaptation in insectivorous subterranean eutherians. The enigma of Mesoscalops montanensis unveiled by geometric morphometrics and finite element analysis

The enigmatic Early Miocene fossorial mammal Mesoscalops montanensis shows one of the most modified humeri among terrestrial mammals. It has been suggested, on qualitative considerations, that this species has no extant homologues for humerus kinematics and that, functionally, the closest extant group is represented by Chrysochloridae. We combine here three dimensional geometric morphometrics, finite element analysis and phylogenetic comparative methods to explore the shape and mechanical stress states of Mesoscalops montanensis as well as of extant and extinct Talpidae and Chrysochloridae under realistic digging simulations. Evolutionary convergence analyses reveal that the shape of Mesoscalops montanensis represents a unique morphology in the context of fossorial mammals and that its functional performance, albeit superficially similar to that of extant Chrysochloridae, still represents a nonconvergent optimum for adaptation to digging. J. Morphol. 276:1157–1171, 2015.

Origin and evolution of Western European moles (genus Talpa, Insectivora): a multidisciplinary approach

ABSTRACT The European representatives of the subfamily Talpinae belong to the monophyletic genus Talpa. Five out of the nine species of Talpa occur in Western Europe. A wide central area is occupied by Talpa europaea, while the small sized Talpa caeca occurs in southern Europe. Three endemic species are restricted to the peripheral areas of the genus range and show a parapatric distribution with respect to T. europaea, i.e. the Iberian T. occidentalis, the southern Italian T. romana and the Balkan T. stankovici. The karyotypes of moles are very conservative, with the majority of the species showing 2n = 34. Allozyme data first allowed to assess the specific status of endemic taxa and the low levels of heterozygosity. Neis genetic distances suggest that T. occidentalis, T. romana and T. stankovici early diverged from an europaea–caeca line. Preliminary results from mtDNA analyses strongly support the monophyly of Western European moles, but are still not able to solve the relationships within this clade. Estimates of time of divergences indicate a basal split of an Eastern and a Western lineage during the Miocene-Pliocene transition, while the divergence among the Western European moles should have occurred in association to the Pliocene-Pleistocene climatic oscillations.