exerted by a struggling prey. The bite force of Albertosaurus was less formidable, however, with the maximum force, by the hind teeth, reaching 3,413 Newtons. Above the eyes were short bony crests that may have been brightly coloured in life and used in courtship to attract a mate. William Abler observed in 2001 that Albertosaurus tooth serrations resemble a crack in the tooth ending in a round void called an ampulla. Tyrannosaurid teeth were used as holdfasts for pulling flesh off a body, so when a tyrannosaur pulled back on a piece of meat, the tension could cause a purely crack-like serration to spread through the tooth. However, the presence of the ampulla distributed these forces over a larger surface area, and lessened the risk of damage to the tooth under strain. The presence of incisions ending in voids has parallels in human engineering. Guitar makers use incisions ending in voids to, as Abler describes, "impart alternating regions of flexibility and rigidity" to wood they work. The use of a drill to create an "ampulla" of sorts and prevent the propagation of cracks through material is also used to protect aircraft surfaces. Abler demonstrated that a plexiglass bar with incisions called "kerfs" and drilled holes was more than 25% stronger than one with only regularly placed incisions. Unlike tyrannosaurs, ancient predators like phytosaurs and Dimetrodon had no adaptations to prevent the crack-like serrations of their teeth from spreading when subjected to the forces of feeding. Classification and systematics Albertosaurus is a member of the theropod family Tyrannosauridae, in the subfamily Albertosaurinae. Its closest relative is the slightly older Gorgosaurus libratus (sometimes called Albertosaurus libratus; see below). These two species are the only described albertosaurines; other undescribed species may exist. Thomas Holtz found Appalachiosaurus to be an albertosaurine in 2004, but his more recent unpublished work locates it just outside Tyrannosauridae, in agreement with other authors. The other major subfamily of tyrannosaurids is the Tyrannosaurinae, which includes Daspletosaurus, Tarbosaurus and Tyrannosaurus. Compared with these robust tyrannosaurines, albertosaurines had slender builds, with proportionately smaller skulls and longer bones of the lower leg (tibia) and feet (metatarsals and phalanges). Below is the cladogram of the Tyrannosauridae based on the phylogenetic analysis conducted by Loewen et al. in 2013. Palaeobiology Growth pattern Most age categories of Albertosaurus are represented in the fossil record. Using bone histology, the age of an individual animal at the