title: Evolution of embryonic developmental period in the marine bird families Alcidae and Spheniscidae: roles for nutrition and predation?
abstract: BACKGROUND
  Nutrition and predation have been considered two primary agents of selection important in the evolution of avian life history traits. The relative importance of these natural selective forces in the evolution of avian embryonic developmental period (EDP) remain poorly resolved, perhaps in part because research has tended to focus on a single, high taxonomic-level group of birds: Order Passeriformes. The marine bird families Alcidae (auks) and Spheniscidae (penguins) exhibit marked variation in EDP, as well as behavioural and ecological traits ultimately linked to EDP. Therefore, auks and penguins provide a unique opportunity to assess the natural selective basis of variation in a key life-history trait at a low taxonomic-level. We used phylogenetic comparative methods to investigate the relative importance of behavioural and ecological factors related to nutrition and predation in the evolution of avian EDP.
  
  
  RESULTS
  Three behavioural and ecological variables related to nutrition and predation risk (i.e., clutch size, activity pattern, and nesting habits) were significant predictors of residual variation in auk and penguin EDP based on models predicting EDP from egg mass. Species with larger clutch sizes, diurnal activity patterns, and open nests had significantly shorter EDPs. Further, EDP was found to be longer among birds which forage in distant offshore waters, relative to those that foraged in near shore waters, in line with our predictions, but not significantly so.
  
  
  CONCLUSIONS
  Current debate has emphasized predation as the primary agent of selection driving avian life history diversification. Our results suggest that both nutrition and predation have been important selective forces in the evolution of auk and penguin EDP, and highlight the importance of considering these questions at lower taxonomic scales. We suggest that further comparative studies on lower taxonomic-level groups will continue to constructively inform the debate on evolutionary determinants of avian EDP, as well as other life history parameters.
authors.affiliation: Department of Biological Sciences, Simon Fraser University, Centre for Wildlife Ecology, Burnaby, British Columbia V5A 1S6, Canada. mark.hipfner@ec.gc.ca
authors.forename: J Mark
authors.initials: JM
authors.lastname: Hipfner
authors.forename: Kristen B
authors.initials: KB
authors.lastname: Gorman
authors.forename: Rutger A
authors.initials: RA
authors.lastname: Vos
authors.forename: Jeffrey B
authors.initials: JB
authors.lastname: Joy
mesh_terms: D000818:Animals; D001499:Bayes Theorem; D005075:Biological Evolution; D046091:Charadriiformes / Q000196:embryology* / Q000235:genetics; D052284:Clutch Size; D004272:DNA, Mitochondrial / Q000235:genetics; D016013:Likelihood Functions; D008954:Models, Biological; D009425:Nesting Behavior; D009747:Nutritional Physiological Phenomena; D010802:Phylogeny; D011235:Predatory Behavior; D017422:Sequence Analysis, DNA; D046429:Spheniscidae / Q000196:embryology* / Q000235:genetics
chemical_list: D004272:DNA, Mitochondrial