Breeding at higher latitude is associated with higher photoperiodic threshold and delayed reproductive development in a songbird.

Many seasonally breeding animals exhibit a threshold day length (critical photoperiod; CPP) for gonadal growth, and populations breeding at higher latitudes typically have a higher CPP. Much less is known about latitudinal variation in CPP in migratory population that winter away from their breeding range and must time their reproduction to match favorable conditions at their destination. To address the relationship between migration, breeding latitude, and CPP, we held two closely related songbird populations in a common environment. One population is resident (Junco hyemalis carolinensis), the other winters in sympatry with the residents but migrates north to breed (Junco hyemalis hyemalis). We gradually increased photoperiod and measured indices of readiness to migrate (fat score, body mass) and breed (cloacal protuberance volume, baseline testosterone, and gonadotropin releasing hormone challenged testosterone). To estimate breeding latitude, we measured hydrogen isotopes in feathers grown the preceding year. As we predicted, we found a higher CPP in migrants than residents, and a higher CPP among migrants deriving from higher as opposed to lower latitudes. Migrants also terminated breeding earlier than residents, indicating a shorter breeding season. To our knowledge, this is a first demonstration of latitudinal variation in CPP-dependent reproductive timing in bird populations that co-exist in the non-breeding season but breed at different latitudes. We conclude that bird populations appear to exhibit local adaptation in reproductive timing by relying on differential CPP response that is predictive of future conditions on the breeding ground.

Fossils reveal the complex evolutionary history of the mammalian regionalized spine.

A unique characteristic of mammals is a vertebral column with anatomically distinct regions, but when and how this trait evolved remains unknown. We reconstructed vertebral regions and their morphological disparity in the extinct forerunners of mammals, the nonmammalian synapsids, to elucidate the evolution of mammalian axial differentiation. Mapping patterns of regionalization and disparity (heterogeneity) across amniotes reveals that both traits increased during synapsid evolution. However, the onset of regionalization predates increased heterogeneity. On the basis of inferred homology patterns, we propose a "pectoral-first" hypothesis for region acquisition, whereby evolutionary shifts in forelimb function in nonmammalian therapsids drove increasing vertebral modularity prior to differentiation of the vertebral column for specialized functions in mammals.

The macroevolutionary consequences of phenotypic integration: from development to deep time.

Phenotypic integration is a pervasive characteristic of organisms. Numerous analyses have demonstrated that patterns of phenotypic integration are conserved across large clades, but that significant variation also exists. For example, heterochronic shifts related to different mammalian reproductive strategies are reflected in postcranial skeletal integration and in coordination of bone ossification. Phenotypic integration and modularity have been hypothesized to shape morphological evolution, and we extended simulations to confirm that trait integration can influence both the trajectory and magnitude of response to selection. We further demonstrate that phenotypic integration can produce both more and less disparate organisms than would be expected under random walk models by repartitioning variance in preferred directions. This effect can also be expected to favour homoplasy and convergent evolution. New empirical analyses of the carnivoran cranium show that rates of evolution, in contrast, are not strongly influenced by phenotypic integration and show little relationship to morphological disparity, suggesting that phenotypic integration may shape the direction of evolutionary change, but not necessarily the speed of it. Nonetheless, phenotypic integration is problematic for morphological clocks and should be incorporated more widely into models that seek to accurately reconstruct both trait and organismal evolution.

Shape, variance and integration during craniogenesis: contrasting marsupial and placental mammals.

Studies of morphological integration can provide insight into developmental patterns, even in extinct taxa known only from skeletal remains, thus making them an important tool for studies of evolutionary development. However, interpreting patterns of integration and assessing their significance for organismal evolution requires detailed understanding of the developmental interactions that shape integration and how those interactions change through ontogeny. Thus far, relatively little comparative data have been produced for this important topic, and the data that do exist are overwhelmingly from humans and their close relatives or from laboratory models such as mice. Here, we compare data on shape, variance and integration through postnatal ontogeny for a placental mammal, the least shrew, Cryptotis parva, and a marsupial mammal, the gray short-tailed opossum, Monodelphis domestica. Cranial variance decreased dramatically from early to late ontogeny in Cryptotis, but remained stable through ontogeny in Monodelphis, potentially reflecting functional constraints related to the short gestation and early ossification of oral bones in marsupials. Both Cryptotis and Monodelphis showed significant changes in cranial integration through ontogeny, with a mixture of increased, decreased and stable levels of integration in different cranial regions. Of particular note is that Monodelphis showed an unambiguous decrease in integration of the oral region through ontogeny, potentially relating to their early ossification. Selection at different stages of development may have markedly different effects if patterns of integration change substantially through ontogeny. Our results suggest that high integration of the oral region combined with functional constraints for suckling during early postnatal ontogeny may drive the stagnant variance observed in Monodelphis and potentially other marsupials.

Of mice and mutations: phenotypic effects of the diabetic db/db and ob/ob mutations on the skull and teeth of mice.

AIM: To compare the phenotypic appearance of the skull bones and teeth of wild type C57BL/6J mice with that of diabetic leptin-deficient (ob/ob) and diabetic leptin receptordeficient (db/db) mice used as models for diabetes. STUDY DESIGN AND METHODS: Skulls were extracted from the carcasses of mice belonging to wild-type C57B/6J mice, db/db mice on a C57BLKS/J background, and ob/ob mice on a C57B/6J background. After removal of overlying tissue, the skulls and mandibles were then left to dehydrate and examined for phenotypic variations in structure and wear. RESULTS: Bone surfaces of the skulls of wild type mice had a whiter and smoother surface compared with a yellowish colour with a grainy texture in the two mutant strains. The frontal, parietal and occipital bones were translucent in the two mutant strains. Breakages of the zygomatic arches and mandibles were more common in the ob/ob and db/db mice than in the wild type mice. Half of the teeth of the db/db mice and 90% teeth of the ob/ob mice showed considerable wear compared with marginal wear in the wild type mice. CONCLUSIONS: These observations suggested that the teeth of the two diabetic mutant strains are exhibiting considerable signs of hypomineralization with increased fragility and decreased bone thickness.

Evolutionary acceleration in the most endangered mammal of Canada: speciation and divergence in the Vancouver Island marmot (Rodentia, Sciuridae).

The Vancouver Island marmot is the most endangered mammal of Canada. Factors which have brought this population to the verge of extinction have not yet been fully elucidated, but the effects of deforestation and habitat fragmentation on survival rates, as well as those of variation in rainfall, temperature, snowpack depth and snowmelt strongly suggest that marmots on the island are struggling to keep pace with environmental changes. Genetic analyses, however, seem to indicate that the Vancouver Island marmot may merely represent a melanistic population of its parental species on the mainland. Were it not for its black pelage colour, it is unlikely that it would have attracted much attention as a conservation priority. Our study uses three-dimensional coordinates of cranial landmarks to further assess phenotypic differentiation of the Vancouver Island marmot. A pattern of strong interspecific divergence and low intraspecific variation was found which is consistent with aspects of drift-driven models of speciation. However, the magnitude of shape differences relative to the putatively neutral substitutions in synonymous sites of cytochrome b is too large for being compatible with a simple neutral model. A combination of bottlenecks and selective pressures due to natural and human-induced changes in the environment may offer a parsimonious explanation for the large phenotypic differentiation observed in the species. Our study exemplifies the usefulness of a multidisciplinary approach to the study of biological diversity for a better understanding of evolutionary models and to discover aspects of diversity that may be undetected by using only a few genetic markers to characterize population divergence and uniqueness.

Paleontology and the comparative method: ancestral node reconstructions versus observed node values.

Comparative methods are used to reconstruct ancestral node values for continuously varying traits. The confidence intervals (CIs) around such estimates may be wider than the range of tip data from which they are calculated. Without historical data with which to compare estimates, it is not clear whether such broad CIs reflect evolutionary lability or methodological imprecision. In this study, a fully resolved phylogeny of fossil carnivorans, in which observed samples are found not only at the tree tips but also along branches and at nodes, is used to compare observed ancestral node values with node estimates based on a Brownian motion model of evolution. As in previous studies, the CIs surrounding node estimates were wider than the range of tree tip values, but observed values fell well within them, reasonably close to the values predicted by comparative methods. Confidence intervals calculated using paleontological rate estimates were comparable to those calculated using only terminal taxa. This implies that evolution of at least some traits is conservative enough for node reconstruction techniques to be useful, despite their large standard errors. The Brownian motion model of evolutionary change was a good predictor of node values.

On morphological clocks and paleophylogeography: towards a timescale for Sorex hybrid zones.

Phylogeography--the study of within-species phylogenetic and geographic divergence--has been primarily the domain of molecular evolutionists because molecular markers record population structure on smaller scales than do traditional morphological traits. But when geometric morphometrics are combined with distance-based phylogenetics molar shape divergence appears to record population-level phylogeny, a fact that allows extant and fossil populations to be combined in a single phylogeographic study. The European Sorex araneus complex--a genetically complicated group composed of multiple karyotypic races and species--illustrates the principle. The phylogeographic patterns revealed by molar shape broadly agree with scenarios based on molecular data and circumstantial evidence. Importantly, the inclusion of fossil samples of known age allows minimum divergence times to be inferred. Some races of S. araneus may have diverged more than 120,000 years ago, but others may have diverged less than 14,000. Supporting evidence that molar shape can be used to reconstruct phylogeographic relationships comes from strong correlations between molar shape distances and both phylogenetic divergence time and cytochrome b sequence divergence in datasets where these variables are known independently (fossil carnivorans from a well-constrained stratigraphic setting and shrew species of the genus Sorex, respectively). However, molar shape may have a 'saturation point' beyond which it is not applicable.