The Apennine brown bear subspecies together with the polar and the Asiatic black bear showed rates of correct classification higher than 80% (Table 3). Shape variation along discriminant function axes was visualized by regressing discriminant function scores on shape variables with the software tpsRegr v. 1.34 (Rohlf, 2015). In Western Europe, the remaining populations of the Cantabrian Mountains, Pyrenees, Eastern Alps and Apennines are extremely small, isolated and seriously endangered (Zedrosser, Dahale, Swenson, & Gerstl, 2001). Implantation of the fertilized egg is delayed, occurring in November or December.  The pre-copulatory mechanisms, including contest and scramble competition, are conditional mating tactics that are used based on an individual's phenotype. In the genus Ursus (n = 56), the correlation between mandible shape and climate persisted with the first pair of axes explaining 35% of covariation (Figure 7c). Moderator:networks12. In addition, the recently developed Kmultiv statistic (Adams, 2014) was quantified based on the interspecific dataset of mandible shape (N = 8) using the package geomorph (Adams & Otárola‐Castillo, 2013). For females, the amount of fat stored before winter is linked with reproductive success: fatter females typically have more and bigger young than do leaner females. Deformation grids at the onset of each axes show relative shape changes from the mean described by each RW vector; (b) the first two discriminant function vectors obtained from a combination of shape coordinates and size. To test for the impact of species and subspecies (for the brown bear only) differences, the sample of Ursus specimens was reduced from 119 (as showed in the RWA) to 114 because subspecies classification was not available for five specimens of U. arctos. Aim of this approach was to test for the impact of climate on taxonomic differentiation at both inter‐ and intrageneric scale (Bubadué et al., 2016; Cáceres et al., 2014; Meloro et al., 2014a,b). U. arctos and U. maritimus were classified with 92.3% and 90.9% of accuracy, respectively, while the Asiatic black bear and American black bear had lower rates (85.2% and 75.0%, respectively). Black bears that are actually brown in colour are most common in western North America.
A Mantel test between the distance matrices obtained from the molecular phylogeny and the shape coordinates results in a relatively high but non‐significant correlation between the two (r = .624; p = .058 after 99,999 permutations). Opposite to the Apennine bear, the mandible of Isabelline bear is slender under the canine region although retains thick corpus in the posterior crushing area. Fruits dominate the diet in summer, and both fruit and mast, especially acorns and beechnuts, constitute most of the fall diet. Cranial distinctiveness in the Apennine brown bear: Genetic drift effect or ecophenotypic adaptation? This does not correspond to a mandible capable of producing relatively high bite force as in other specialized predatory carnivores (Christiansen, 2007), but we note a more slender profile indicative of soft food consumption being this species a specialized hunter of marine mammals, characterized by high percentage of fat in their body tissues (Stirling & Archibald, 1977; van Heteren et al., 2016). Phylogenetic relationships within Ursus arctos and Ursus maritimus, and amplified product length polymorphism (APLP) band patterns of U. arctos mtDNA haplogroups. More work is needed to clarify bear taxonomy, but our analyses strongly support the validity of Apennine and Isabelline brown bear subspecies thus challenging future conservation efforts. Carnivora (continued from Volume I), Suborders Aeluroidea (part) and Arctoidea, Geographic variation in size in North American brown bears, Ursus arctos L., as indicated by condylobasal length, Ecology of the Asiatic black bear (Ursus thibetanus) in Sichuan, China, Statistical power comparisons among alternative morphometric methods, Use of two‐block partial least‐squares to study covariation in shape, Extensions of the Procrustes method for the optimal superimposition of landmarks, Ecomorphological indicators of feeding behaviour in the bears (Carnivora: Ursidae), Biomechanical consequences of rapid evolution in the polar bear lineage, Aspects of predation of seals by polar bears, Rapid radiation events in the family ursidae indicated by likelihood phylogenetic estimation from multiple fragments of mtDNA, Mitochondrial DNA phylogeography of the North American brown bear and implications for conservation, Mammal species of the world: a taxonomic and geographic reference, Sexual dimorphism and geographic variation in the skull of the Ezo Brown Bear (Ursus arctos yesoensis), Status and management of the brown bear in Europe, Geometric morphometrics for biologists: A primer. Our mandible data support separation between the two genera and also fail to identify any possible convergence between Melursus and Tremarctos previously proposed by Kitchener (2010).
Size and shape data were collected on 169 mandibles belonging to the eight extant species from different areas of their range (Appendix 1, Table S1). Bears are distinguished from other carnivores by their size, stout body shape, round ears, long snout, and short tail. This happens in all cases (Figure 7a, b, c) and explains the lack of a significant difference in the relatively large (60 degrees) angle vector between Ursus taxa (Figure 7c) and the other non‐hibernating species (Figure 7b). Variation partitioning allows testing for contribution of climate, size, sex, and taxonomic categorization on mandibular shape variance, taking their interaction into account. ; Wilk's lambda = 0.001; chi‐square = 1147.58, df = 114; p < .0001), a strong deformation in the molar area and a higher ramus distinguished the panda and the Malayan bear from the other taxa. Guidarelli research visit to Denmark was financially supported by FP7 Synthesys project (DK‐TAF‐5104): Morphological variation in the mandible of Delphinidae. Breeding begins in the spring and peaks during June and July. Ursidae: Tremarctinae) from North America, and the cave bear (Ursus spelaeus Rosenmüller & Heinroth, Ursidae: Ursinae) from Europe were among the most massive mammalian carnivorans ever to have lived.
van Heteren et al.
vs RW2 (17.31% var.) The climatic variables that mostly showed a correlation with non‐Ursus bear mandible shape were Bio3, 4, 5, 7, 12, 14, and 15. ; Wilk's lambda = 0.006; chi‐square = 516.243, df = 119; p < .0001) better characterized these species with the polar bear occupying negative scores, due to its thin corpus, and the Apennine bear positive ones with a thick and wide corpus on the anterior region. , The mating system is generally characterised by two main components, the search phase and the encounter phase.
A thick corpus related to tropical environments (high temperatures, high precipitation, and lower seasonality) was typical of taxa like the Malayan bear.
The cluster obtained from mandible shape distances showed a very good cophenetic correlation (r = .912). The Apennine bear also differed in size from alascensis and horribilis. This analysis was performed using the R package vegan (Oksanen et al., 2012). The family Ursidae comprises powerful and large terrestrial members of the mammalian order Carnivora that evolved in the northern hemisphere and are currently distributed in Eurasia, North Africa, and the Americas (Herrero, 1999). Missing the third dimension in geometric morphometrics: How to assess if 2D images really are a good proxy for 3D structures? Such characteristics reflect ecological adaptations, but they also allow characterizing bear taxonomy in more details. Indeed, variation partition shows climate to have a non‐significant influence when in isolation on the mandibular shape differences of Ursus taxa and brown bear subspecies (Table 5). Actual gestation then lasts 60–70 days, and one to four young cubs are born in January or February. Deformation grids at the onset of each axes show relative shape changes from the mean described by each RW vector; (b) the first two discriminant function vectors obtained from a combination of shape coordinates and size. To better characterize mandibular shape variation within our sample, a variation partitioning approach was also applied using shape variables as response and climate (continuous standardized variables inclusive of all the 19 bioclim), taxonomy (categorical variable), sex, and size (continuous variable, ln CS) as explanatory variables (Cáceres et al., 2016; Cardini et al., 2007; Meloro et al., 2014a,b). The brown bear was also distinct in size, and it overlapped with the polar bear only (Table S2). They positively correlated to each other (r = .677, p < .002) with negative scores being occupied by the polar bear that is extremely adapted to cold condition (lower annual temperatures, highly seasonal), while on the positive score, the Asiatic black bear occupied warmer areas. The DFA extracted eight vectors of which the first six (98% of tot var.) The scatterplot of RW1 (23.56% var.) van Heteren et al.
 There is a very loose dominance hierarchy within bear mating systems due to their solitary nature. A K close or bigger than 1 demonstrates that phenotypic differences between species followed a pattern expected by Brownian motion of evolution. Amano, Oi, and Hayano (2004) reported significant differences in skull morphologies of two Japanese populations of U. thibetanus and a pattern of geographical changes occur also in our sample (Figure 7c). Previous research mostly focused on the functional link between these traits and diet (Figueirido et al., 2009; van Heteren et al., 2016; Meloro, 2011), even if strong taxonomic signal always emerged in these datasets. For Eurasian brown bears, there are not many morphological studies focusing on subspecies description (Baryshnikov, Mano, & Masuda, 2004; Mihaylov et al., 2013) with few of them using updated morphometric approaches to highlight distinctiveness of the endangered Apennine subspecies, U. arctos marsicanus Altobello, 1921 (Colangelo et al., 2012; Loy, Genov, Galfo, Jacobone, & Vigna Taglianti, 2008). Main differences with molecular phylogeny were due to the position of T. ornatus and the American black bear that, based on mandible shape, appeared to be more similar to Ursus spp. Diet reconstruction in cave bears from craniodental morphology: past evidences, new results and future directions, Journal of Zoological Systematics and Evolutionary Research, https://doi.org/10.2305/iucn.uk.2008.rlts.t41687a10513074.en, https://doi.org/10.2305/iucn.uk.2008.rlts.t41688a10513490.en, National Museum of Natural History, Sofia, Bulgaria, Zoological Museum, University of Copenhagen, Surgara State CP India, Mabaraj Kanar of Bikaner, India, Stuffed specimen in Collect Namerica coll.
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