A molecular genetic time scale demonstrates Cretaceous origins and multiple diversification rate shifts within the order Galliformes (Aves).

The phylogeny of Galliformes (landfowl) has been studied extensively; however, the associated chronologies have been criticized recently due to misplaced or misidentified fossil calibrations. As a consequence, it is unclear whether any crown-group lineages arose in the Cretaceous and survived the Cretaceous-Paleogene (K-Pg; 65.5 Ma) mass extinction. Using Bayesian phylogenetic inference on an alignment spanning 14,539 bp of mitochondrial and nuclear DNA sequence data, four fossil calibrations, and a combination of uncorrelated lognormally distributed relaxed-clock and strict-clock models, we inferred a time-calibrated molecular phylogeny for 225 of the 291 extant Galliform taxa. These analyses suggest that crown Galliformes diversified in the Cretaceous and that three-stem lineages survived the K-Pg mass extinction. Ideally, characterizing the tempo and mode of diversification involves a taxonomically complete phylogenetic hypothesis. We used simple constraint structures to incorporate 66 data-deficient taxa and inferred the first taxon-complete phylogenetic hypothesis for the Galliformes. Diversification analyses conducted on 10,000 timetrees sampled from the posterior distribution of candidate trees show that the evolutionary history of the Galliformes is best explained by a rate-shift model including 1-3 clade-specific increases in diversification rate. We further show that the tempo and mode of diversification in the Galliformes conforms to a three-pulse model, with three-stem lineages arising in the Cretaceous and inter and intrafamilial diversification occurring after the K-Pg mass extinction, in the Paleocene-Eocene (65.5-33.9 Ma) or in association with the Eocene-Oligocene transition (33.9 Ma).

Extreme intraclutch egg-size dimorphism in Eudyptes penguins, an evolutionary response to clutch-size maladaptation.

Eudyptes penguins (six species) are uniquely characterized by a two-egg clutch with extreme intraclutch egg-size dimorphism (ESD): the first-laid A-egg is 17.5%-56.9% smaller than the B-egg. Although A-eggs are viable, they almost never produce fledged chicks (genus average <1%). Using classical life-history theory and phylogenetic comparative methods, we demonstrate a marked slowdown in the life history of Eudyptes: age of first reproduction is 52% later and annual fecundity 48% lower compared with other two-egg clutch penguin species. All six Eudyptes species have retained a two-egg clutch, despite this pronounced life-history slowdown; this suggests evolutionary mismatch between clutch size and chicks fledged per clutch. Consistent with this, we show that Eudyptes fledge 43% fewer chicks per clutch than other two-egg clutch penguin species. Extreme intraclutch ESD in Eudyptes is associated primarily with a uniform (5%) increase in relative B-egg size, and B-egg size has evolved in accord with life history. We further show that intraclutch ESD is positively correlated with age of first reproduction in Eudyptes but not in other two-egg clutch penguin species. We argue that Eudyptes' persistent failure to evolve a one-egg clutch constitutes a unique genus-wide evolutionary maladaptation and that extreme intraclutch ESD evolved as a correlated response to selection favoring a slower life history imposed by their extreme pelagic overwintering and migration ecology.

Digestive organ sizes and enzyme activities of refueling western sandpipers (Calidris mauri): contrasting effects of season and age.

We examined seasonal and age-related variation in digestive organ sizes and enzyme activities in female western sandpipers (Calidris mauri) refueling at a coastal stopover site in southern British Columbia. Adult sandpipers exhibited seasonal variation in pancreatic and intestinal enzyme activities but not in digestive system or organ sizes. Spring migrants had 22% higher total and 67% higher standardized pancreatic lipase activities but 37% lower total pancreatic amylase activity than fall migrants, which suggests that the spring diet was enriched with lipids but low in glycogen. Spring migrants also had 47% higher total intestinal maltase activity as well as 56% higher standardized maltase and 13% higher standardized aminopeptidase-N activities. Spring migrants had higher total enzymic capacity than fall migrants, due primarily to higher total lipase and maltase activities. During fall migration, the juvenile's digestive system was 10% larger than the adult's, and it was composed differently: juveniles had a 16% larger small intestine but a 27% smaller proventriculus. The juvenile's larger digestive system was associated with lower total enzymic capacity than the adult's due to 20% lower total chitinase and 23% lower total lipase activities. These results suggest that juvenile western sandpipers may process food differently from adults and/or have a lower-quality diet.